FAO Fisheries Synopsis No. 125, Volume 4, Part 1FAO SPECIES CATALOGUEFIR/S125 Vol. 4, Part 1VOL. 4, PART 1 SHARKS OF THE WORLDAN ANNOTATED AND ILLUSTRATED CATALOGUEOF SHARK SPECIES KNOWN TO DATEUNITED NATIONS DEVELOPMENT PROGRAMMEFOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONSFAO Fisheries Synopsis- No. 125, Volume 4, Part 1 FIR/S125 Vol. 4, Part 1FAOSPECIES CATALOGUEVOL. 4 SHARKS OF THE WORLDAn Annotated and Illustrated Catalogueof Shark Species Known to DatePart 1 - Hexanchiformes to Lamniformesprepared byLeonard J.V. CompagnoTiburon Center for Environmental StudiesSan Francisco State UniversityTiburon, California 94920, USAPrepared with support from theUnited Nations Development Programme (UNDP)and the American Elasmobranch SocietyUNITED NATIONS DEVELOPMENT PROGRAMMEFOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONSRome 1984PREPARATON OF THS DOCUMENT Thepresentpublication,preparedundertheUNDP/FAOProjectfortheSurveyanddentificationofWorld-MarineFishResources(GLO/82/001),isthefourthworldwidespeciescatalogueissuedwithintheFAOFisheries Synopses series. Workonthiscataloguewas initiatedbytheauthor manyyearsagowiththe preparation ofa simplelist of sharkspeciesincludingonlymostelementaryinformationsuchasscientificsynonymies,geographicaldistributions, sizes,etc.,precededbyanillustratedkeytosharkfamilies.However,withtheevolvementofamoreambitious formatfortheseries,andparalleltothepreparationofthevolumesonscombridsandcephalopods,itbecame necessarytoexpandtheoriginalmanuscriptverysubstantially,ataskwhichtheauthoraccomplishedwithhis usualenthousiasmandthoroughness,inspiteofthedifficultconditionsunderwhichhehadtoworkduringthepast years. Theworkwasfacilitatedbytheauthor'sinvolvementinseveralregionalsetsofFADSpeciesdentification SheetsforFisheryPurposes,e.g.theWesternCentralAtlantic,EasternCentralAtlanticandWesternndian Ocean,butthenumerousgapsininformationonspeciesfromthendo-Pacificregioncouldnothavebeenfilled withouttheauthor'srecentextensivefieldworkinthatarea,whichwasmadepossiblethankstothesupportof several institutions both within and outside the USA, coordinated by the American Elasmobranch Society. n the final stages of the work, the author could count on the generous assistance nstitute of the J.L.B. Smithof chthyology, Grahamstown, South Africa. TheindexesofscientificandcommoninternationalFAOspeciesnamesandoflocalspeciesnameswere prepared in collaboration with FAO's Fishery nformation, Data and Statistics Service. llustrations were adapted and redrawn by a wide variety of sources, especially from Okutani (1980). IIIustrators :P. Last ri co(pri nci pal i l l ust rat or), O. Li donni ci , E. D' Ant oni and P. -L. sol a, FAO, Rome.FAO FamiIy and Species names in French:J.-C.Quro,nstitutScientifiqueet Technique des Pches Maritimes, La Rochelle, France. TechnicaI Editors :W.FischerandC.E.Nauen,FisheryResourcesandEnvironment Division, FAO. ABSTRACT ThisisthefourthintheFAOseriesofworldwideannotatedandillustrated catal oguesofmaj orgroupsoforgani smsthatentermari nefi sheri es.Thepresent vol umeincludes342shark.speciesbelongingto8ordersand31families.tprovidesa comprehensiveandillustratedkeytoallordersandfamiliesofsharks,withaglossary oftechni cal termsandmeasurements.Wi thi neachfami l yaregi veni ndi vi dualaccountsofspecies,whichincludedrawings,scientificandvernacularnames,informa-tiononhabi tat,bi ol ogyandfi sheri es,andadi stri buti onmap;formostfami l i esthere i sal soakeytogenera.Theworkisfullyindexedandthereisamplereferenceto pertinent literature. Distribution: Author FAO Fisheries Department FAO Regional Fisheries Officers Regional Fisheries Councils and For bibliographic purposes this document should be cited as follows: Compagno, L.J.V.,FAO species catalogue. Vol. 4. Sharks of the world. An annotated and illustrated catalogue of sharks species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fish Synop., (125)Vol.4,Pt.1: 249 p. 1984click for next pageTABLE OF CONTENTS CodePage1.INTRODUCTION .................................................................................................................................................... 11.1 Plan of the Catalogue......................................................................................................................................... 21.2 Technical Terms and Measurements................................................................................................................. 82.ORDER HEXANCHIFORMES - FRILLED AND COW SHARKS ............................................................................................ ..132.1 FAMLY CHLAMYDOSELACHDAE - Frilled sharks......................................... CHLAM ....................................................... 14Chlamydoselachus ..................................................................................... CHLAM Chlam ....................................... 14C. anguineus................................................................................................. CHLAM Chlam 1 ...............................142.2 FAMLY HEXANCHDAE - Cowsharks, Sixgill sharks,Sevengill sharks................................... HEX ...................................................... 15Heptranchias .....................................................................................HEX Hept ................................................ 17H. perlo............................................................................................. HEX Hept 1..........................................17Hexanchus .......................................................................................HEX Hex................................................ 19H. griseus ................................................................................................ HEX Hex 1................................................19H. vitulus ................................................................................................ HEX Hex 2................................................. 20Notorynchus ................................................................................................ HEX Not ................................................. 22N. cepedianus............................................................................................ HEX Not 1 .................................................. 223.ORDER SQUALIFORMES - DOGFISH SHARKS ..................................................................................................................243.1 FAMLY ECHNORHNDAE -Bramble sharks ........................................................ECHN ..................................................... 25Echinorhinus ................................................................................................ ECHN Echin ....................................... 25E. brucus ................................................................................................ ECHN Echin 1 ....................................... 26E. cookei ................................................................................................ ECHN Echin 2 ....................................... 273.2 FAMLY SQUALDAE - Dogfish sharks .............................................................. SQUAL.................................................28Aculeola ..................................................................................................... SQUAL Acul........................................... 34A. nigra ................................................................................................ SQUAL Acul 1.......................................... 34Centrophorus ................................................................................................ SQUAL Centr ........................................... 35C. acus................................................................................................ SQUAL Centr 5.......................................... 36C................................................................................................ SQUAL Centr 6......................................... 38C. lusitanicus ...............................................................................SQUAL Centr 2 ......................................39C. moluccensis.............................................................................................. SQUAL Centr 7................................... 40C. niaukang .............................................................................................. SQUAL Centr 8.........................................41C. squamosus............................................................................. SQUAL Centr 3 .................................. 43C. tesselatus................................................................................SQUAL Centr 9.................................... 44C. uyato ..........................................................................................SQUAL Centr 4................................... 45Centroscyllium ....................................................................................SQUAL Centro ....................................46C. fabricii....................................................................................SQUAL Centro 1 .........................................47C. granulatum.................................................................................SQUAL Centro 2 .........................................49C. kamoharai........................................................................................ SQUAL Centro 3.........................................49C. nigrum...................................................................................SQUAL Centro 4.........................................50C. ornatum ..................................................................................SQUAL Centro 5.........................................51C. ritteri ....................................................................................... SQUAL Centro 6.........................................52 - iii -. granulosus ................................................................................................ SQUAL Centr 1......................................... 37C. harrissoniclick for previous page- iv -Centroscymnus................................................................................... !"#$ SQUAL Centros .........................&'($53C. coelclepis.................................................................................. SQUAL Centros .......................... 55C. crepidater...................................................................................SQUAL Centras ............................... 56C. cryptacanthus..................................................................................SQUAL Centros ........................... 57C. macracanthus ..................................................................................SQUAL Centros..........................58C. owstoni ..................................................................................SQUAL Centros ..........................59C. plunketi ......................................................................................SQUAL Centros ..........................60Cirrhigaleus ......................................................................................... SQUAL Cirrh ..............................61 C. barbifer ..........................................................................................SQUAL Cirrh 1.............................61Dalatias ..............................................................................................D. licha..........................................................................................SQUAL Dal...............................SQUAL Dal 1 .............................6363Deania .......................................................................................................... SQUAL Dean .............................. 65D. calcea ................................................................................................... SQUAL Dean 1..........................65D. histricosa ................................................................................... SQUAL Dean 3 .......................... 66D. profundorum ..................................................................................... SQUAL Dean 4..........................67D. quadrispinosum .................................................................................. SQUAL Dean 2..........................68Etmopterus ............................................................................................. SQUAL Etmo.............................69E. baxteri ....................................................................................................................SQUAL Etmo 5..........................71E. brachyurus.............................................................................................. SQUAL Etmo 6 ..........................72E. bullisi .......................................................................................................SQUAL Etmo 7 ..........................74E.decacuspidatus ..................................................................................SQUAL Etmo 8 ..........................75E. gracilispinis ..................................................................................SQUAL Etmo 9 ..........................76E. granulosus ..................................................................................SQUAL Etmo 10 .......................... 77E. hillianus .......................................................................................SQUAL Etmo 11........................... 78E. lucifer .....................................................................................................SQUAL Etmo 12 .......................... 79E. polli ......................................................................................................SQUAL Etmo 1 ..........................80E. princeps ....................................................................................... SQUAL Etmo 4 ...........................81E. pusillus ........................................................................................ SQUAL Etmo 3 ..........................82E. schultzi......................................................................................SQUAL Etmo 13 ..........................83E. sentosus ....................................................................................... SQUAL Etmo 14 ..........................84E. spinax ............................................................................................. SQUAL Etmo 2 ..........................85E......................................................................................... unicolorSQUAL Etmo 15 ..........................86E. villosus ........................................................................................... SQUAL Etmo 16 ..........................87E. virens ............................................................................................ SQUAL Etmo 17 ..........................88Euprotomicroides ........................................................................................ SQUAL Eup .................................89E. zantedeschia ........................................................................................ SQUAL Eup 1..................................89Euprotomicrus ........................................................................................SQUAL Eupr ................................. 90E. bispinatus ........................................................................................SQUAL Eupr 1............................90Heteroscymnoides ........................................................................................SQUAL Hetero................................ 91H. marleyi........................................................................................SQUAL Hetero 1.............................. 92sistius ....................................................................................................SQUAL sist ................................... 93. brasiliensis ........................................................................................SQUAL sist 1................................ 93. plutodus........................................................................................SQUAL sist 2................................ 95Scymnodalatias ........................................................................................SQUAL Scym.............................96S. sherwoodi ........................................................................................SQUAL Scym 1 ..............................97Scymnodon ............................................................................................. SQUAL Scymn.............................97S. obscurus..................................................................................................SQUAL Scymn 2..........................98S. ringens .............................................................................................SQUAL Scymn 1 .......................... 99S. squamulosus ........................................................................................SQUAL Scymn 3 .......................... 101 CodePageSomniosus ........................................................................................................ SQUAL Somn .............................. 102S. microcephalus ..............................................................................SQUAL Somn 2..............................103S. pacificus ............................................................................................SQUAL Somn 3..............................105S. rostratus ..............................................................................................SQUAL Somn 1..............................106Squaliolus..........................................................................................SQUAL Squa...............................108S. laticaudus .................................................................................................. SQUAL Squa 1..............................108Squalus................................................................................................ SQUAL Squal ..............................109S. acanthias .............................................................................................. SQUAL Squal 1...........................111S. asper ........................................................................................................ SQUAL Squal 5............................114S. blainvillei .....................................................................................SQUAL Squal 3 ..............................115S................................................................................... . cubensisSQUAL Squal 2 ..............................116SQUAL Squal 7 ..............................117S. megalops.................................................................................SQUAL Squal 4 ..............................118S. melanurus ................................................................................. SQUAL Squal 8 ..............................120S. mitsukurii ................................................................................... SQUAL Squal 6..............................121S. rancureli .................................................................................... SQUAL Squal 9..............................1223.3 FAMLY OXYNOTDAE - Rough sharks .......................................................... OXYN......................................... 124Oxynotus ............................................................................................ OXN............................... Oxyn124O. bruniensis...........................................................................................OXYN Oxyn 3.............................. 125O. caribbaeus...........................................................................................OXYN Oxyn 1.............................. 126O. centrina ...........................................................................................OXYN Oxyn 2..............................127O. paradoxus........................................................................................... OXYN Oxyn 4 ..............................1284.ORDER PRISTIOPHORIFOMES - SAW SHARKS...........................................................................................1304.1 FAMLY PRSTOPHORDAE - Saw sharks..................................................... PRSTOP..................................130Pliotrema...........................................................................................PRSTOP Plio ..............................132P. warreni ...........................................................................................PRSTOP Piio 1.............................. 132Pristiophorus ...........................................................................................PRSTOP Prist............................. 133P. cirratus ...........................................................................................PRSTOP Prist 2........................... 133P. japonicus...........................................................................................PRSTOP Prist 3........................... 134P. nudipinnis...........................................................................................PRSTOP Prist 4.......................... 135P. schroederi...........................................................................................PRSTOP Prist 1............................ 1365.ORDER SQUATINIFORMES - ANGELSHARKS.................................................................................................1385.1 FAMLY SQUATNDAE - Angelsharks, Sand devils ...................................... SQUAT.....................................138 Squatina..........................................................................................SQUAT Squat ..............................138S. aculeata ...........................................................................................SQUAT Squat 4..............................140S. africana...........................................................................................SQUAT Squat 5 ..............................141S. argentina .................................................................................... SQUAT Squat 6 ..............................142S. australis...........................................................................................SQUAT Squat 7..............................143S. californica ................................................................................... SQUAT Squat 8 ..............................144S. dumeril ...........................................................................................SQUAT Squat 2..............................145S. formosa...........................................................................................SQUAT Squat 9..............................146S. japonica...........................................................................................SQUAT Squat 10..............................147S. nebulosa...........................................................................................SQUAT Squat 11..............................148S. oculata...........................................................................................SQUAT Squat 3..............................149S. squatina ...........................................................................................SQUAT Squat 1..............................150S. tergocellata ...........................................................................................SQUAT Squat 12..............................151S. tergocellatoides ........................................................................................... SQUAT Squat 13..............................152 - v -S. japonicus.................................................................................- vi - CodePage6.ORDER HETERODONTIFORMES - BULLHEAD SHARKS...........................................................................1546.1FAMLY HETERODONTDAE - Bullhead sharks, Horn sharks.............. HET........................................154Heterodontus.......................................................................................HET Het .................................155H. francisci .......................................................................................... HET Het 2 ..............................156H.galeatus .......................................................................................... HET Het 3 ..............................157H. japonicus .......................................................................................... HET Het 4 ..............................158H. mexicanus .......................................................................................... HET Het 5 ..............................159H.portusjacksoni...................................................................................... HET Het 6 ..............................160H. quoyi ......................................................................................................... HET Het 7 ..............................162H. ramalheira .......................................................................................... HET Het 1 ..............................163 H.zebra ............................................................................................HET Het 8 ..............................1647.ORDER ORECTOLOBIFORMES - CARPET SHARKS ................................................................................1657.1FAMLY PARASCYLLDAE - Collared carpet sharks ..............................PARASC..............................166 Cirroscyllium ................................................................................................PARASC Cirr.......................167C. expolitum ..........................................................................................PARASC Cirr 1 .........................168C. formosanum..........................................................................................PARASC Cirr 2 ..........................168C. japonicum..........................................................................................PARASC Cirr 3 ............................ 169 Parascyllium ....................................................................................................PARASC Parasc ........................ 170P. collare..........................................................................................PARASC Parasc 1 ...................170P. ferrugineum .......................................................................................... PARASC Parasc 2 ....................... 171P. multimaculatum ..........................................................................................PARASC Parasc 3 ........................ 172P. variolatum..........................................................................................PARASC Parasc 4 ...................... 1737.2FAMLY BRACHAELURDAE - Blind sharks........................................... BRACH..........................................173 Brachaelurus .............................................................................................. BRACH Brach .............................. 175B. waddi ............................................................................................. BRACH Brach 1 .............................. 175 Heteroscyllium .......................................................................................... BRACH Hete .............................. 176H. colcloughi..........................................................................................BRACH Hete 1.............................. 1767.3FAMLY ORECTOLOBDAE - Wobbegons ................................................ORE...................................................177 Eucrossorhinus ..........................................................................................ORE Eucro ......................................178E. dasypogon..........................................................................................ORE Eucro 1 ..................................179 Orectolobus ..........................................................................................ORE Ore....................................180O. japonicus ..........................................................................................ORE Ore 1..................................... 181O. maculatus..........................................................................................ORE Ore 2.................................. 182O. ornatus .......................................................................................... ORE Ore 3 ..................................184O. wardi ............................................................................................................ORE Ore 4.................................185 Sutorectus...............................................................................................ORE Sut ....................................186S. tentaculatus ..........................................................................................ORE Sut 1 ..................................1877.4FAMLY HEMSCYLLDAE - Bamboo sharks ....................................... HEMS...................................188 Chiloscyllium ..............................................................................................HEMS Chilo ..............................189C. arabicum ........................................................................................HEMS Chilo 3 ..............................190C. caerulopunctatum ................................................................................HEMS Chilo 4 ..............................190C. griseum ..........................................................................................HEMS Chilo 1..............................191C. indicum..........................................................................................HEMS Chilo 2..............................193C. plagiosum .................................................................................. HEMS Chilo 5..............................194C. punctatum ......................................................................................HEMS Chilo 6 ..............................194 8. Hemiscyilium.............................. ............................................................ Code HEMS Hemis..............................Page195 H. freycineti ..........................................................................................HEMS Hemis 1 .............................. 196H. hallstrorni.......................................................................................... HEMS Hemis 2 .............................. 197H. ocellatum.......................................................................................... HEMS Hemis 3 .............................. 197H. strahani.......................................................................................... HEMS Hemis 4 .............................. 198H. trispeculare.......................................................................................... HEMS Hemis 5 .............................. 1987.5FAMLY STEGOSTOMATDAE - Zebra sharks...................................STEG.................................200 Stegostoma..........................................................................................STEG Steg..................................200S. fasciatum..........................................................................................STEG Steg 1...................................2007.6FAMLY GNGLYMOSTOMATDAE - Nurse sharks ..............................GNGL.......................................202Ginglymostoma ..........................................................................................GNGL Gingl..............................204G. brevicaudatum ..........................................................................................GNGL Gingl 2 .............................. 204C. cirratum ..........................................................................................GNGL Gingl 1 .............................. 205Nebrius ................................................................................................GNGL Neb .............................. 207N. ferrugineus .....................................................................................GNGL Neb 1 .............................. 2077.7FAMLY RHNODONTDAE - Whale sharks ..........................................RHN.........................................209 Rhiniodon ....................................................................................................RHN Rhin .............................. 209R. typus .......................................................................................... RHN Rhin 1..............................210ORDER LAMNIFORMES - MACKEREL SHARKS ............................................................................................... 2128.1FAMLY ODONTASPDDAE - Sand tiger sharks .................................... ODONT.................................... 214 Eugomphodus............................................................................................ODONT Eug..............................215E. taurus .................................................................................................ODONT Eug 1..............................216E. tricuspidatus ..........................................................................................ODONT Eug 2..............................218 Odontaspis .......................................................................................... ODONT Odont..............................219O. ferox..........................................................................................ODONT Odont 1..............................219O. noronhai ..........................................................................................ODONT Odont 2..............................2218.2FAMLY MTSUKURNDAE - Goblin sharks ..............................................MTSU.......................................222 Mitsukurina .......................................................................................... MTSU Mitsu..............................222M. owstoni .......................................................................................... MTSU Mitsu 1..............................2238.3FAMLY PSEUDOCARCHARDAE - Crocodile sharks .............................. PSEUD..................................... 224 Pseudocarcharias .......................................................................................... PSEUD Pseud .............................. 225P. kamoharai .......................................................................................... PSEUD Pseud 1 .............................. 2258.4FAMLY MEGACHASMDAE - Megamouth sharks ...............................MEGA.......................................227 Megachasma..........................................................................................MEGA Mega..............................227M. pelagios ..........................................................................................MEGA Mega 3.............................. 2278.5FAMLY ALOPDAE - Thresher sharks ............................................................ALOP........................................228 Alopias .... .............................................................................................ALOP Atop .............................. 229A. pelagicus ..........................................................................................ALOP Atop 3 .............................. 229A. superciliosus ..........................................................................................ALOP Atop 1...............................231A. vulpinus ..........................................................................................ALOP Atop 2 .............................. 232 - vii -!"#$&'($8.6FAMLY CETORHNDAE - Basking sharks ..................................................CETOR .........................................233 Cetorhinus ..........................................................................................CETOR Cetor..............................233C. Maximus ..........................................................................................CETOR Cetor 1..............................2348.7FAMLY LAMNDAE - Mackerel sharks, Porbeagles, White sharks....LAMN.............................................. 237 Carcharodon..........................................................................................LAMN Car .................................238C. carcharias.......................................................................................... LAMN Car 1................................238 surus........................................................................................................................ LAMN sur.................................242. oxyrinchus.......................................................................................... LAMN sur 1..............................242. paucus ..........................................................................................LAMN sur 2..............................244 Lamna ..................................................................................................LAMN Lamn..............................246L. ditropis .......................................................................................... LAMN Lamn 2 .............................. 246L. nasus .......................................................................................... LAMN Lamn 1.............................. 247 - viii -click for next page 1. INTRODUCTION Thiscatalogueincludesallthedescribedspeciesoflivingsharksandtheirsynonyms,includingspeciesof considerableandmajorimportancetofisheriesaswellasthoseofpotential,limited,andnocurrentuse.The catalogue fills a need for a comprehensive review of sharks of the world in a form accessible to fisheries workers aswellassharkbiologists,peoplewhoencountersharksduringthecourseofworkinthesea,andtheinterested public.nrecentyearstherehasbeenamarkedincreaseinourknowledgeofsharksystematics;andformerly difficult, poorly known groups of species have yielded to revisionary work. However, with a single exception there hasbeennocomprehensiveworkinthepast70yearslistingallsharkspeciesandtheirsynonyms.Earlypost-LinnaeanworkersfollowedthetraditionofLinnaeus'(1758)SystemaNaturaeinattempting,tolist,characterize and classify all known living sharks, but these workerswere hampered by the Linnaean system,which allowed only asinglegenus,Squalus,forsharks.Someofthemostimportantearlycomprehensiveworksarethoseof Bonnaterre(1788),Gmelin(1789),Bloch&Schneider(1801),andCuvier(1817,1829).Bythebeginningofthe nineteenthcenturytheLinneanSqualuswasundergoingfragmentation,withtheworksofRafinesque(1810), Blainville(1816)andCuvier(1817introducingthemostinnovationsofthissortpriortoMller&Henle's revisions. TheadventofMller&Henle'sepocalSystematischeBeschreibungderPlagiostomen(Mller&Henle,1838-1841)essentiallyplacedtheclassificationofsharksandtheircloserelatives,theraysorbatoids)onamodern footing.ThePlagiostomenisacomprehensivereviewandsynthesisoftheworkofpreviouswritersandabold stepbeyondthechaosofthepreviouscentury.tdividedtheelasmobranchfishes(plagiostomes,orsharksand rays)intomanyfamiliesandgenera,mostofwhicharerecognizedtoday.Ofthesharks,some13of(Mller& Henle'sgeneraareincurrentuse.ThePlagiostomenprobablyisthemostimportantsingleworkthatbroadly covers the systematics of sharks and rays. Subsequentcomprehensivereviews,includingthoseofGray(1851),Dumeril(1865),Gnther(1870),and Garman(1913)followedtheconventionsofMller&Henlewithconsiderablemodifications.Gill;1862,1873, 1896)reviewedthegeneraandclassificationofsharkswhileEngelhardt(1913)presentedaconcisechecklistof livingsharksalongwithareviewoftheirzoogeography.SincetheworksofGarmanandEngelhardttherehave beenpartialandregionalreviewsofthe sharksofconsiderable importance,including Fowler(1929,1941),White (1937),Whitley(1940,revisedasWhitley&Pollard,1980),Bigelow&Schroeder(1948),Garrick&Schultz(1963), and more recently Springer (1966, 1979), Garrick (1967, 1967a, 1979), Compagno (1970, 1973a,c, 1979, .1982), Bass, D'Aubrey & Kistnasamy (1973, 1975c, 1976), Heemstra (1973), Nakaya (1975), Applegate et al. 1981, and Cadenat & Blache (1981). Steuben & Krefft (1978) is a semipopular work listing many species of sharks. Shiino (1976) and Lindberg,Heard&Rass(1980)havecompiledlistsofvernacularnamesofworldfishes,whichincludemanyshark species. TheonlymoderncomprehensiveworklistingthelivingsharksiscontainedinFowler'spartiallypublished "CatalogofWorldFishes"(sharksinFowler,1966-1969).Thiswaspublishedposthumouslyandisderivedfrom Fowler'simmensecardcatalogueonworldfishes,whichisapparentlythemostvoluminousandcomprehensive databaseofitskindinexistenceapartfromthePiscessectionsoftheZoologicalRecord.Unfortunately,thesharksectionoftheCataloghasmanyerrorsandshowsthedifficultiesthatarisewhenacompilationofspeciesis madewithout the necessary revisionarywork on many of the groups compiled. Fowler's catalogue of sharks isalso outdatedbyrevisionaryworksubsequenttoitslastentries(datedat1958).Althoughaninvaluablesourcework, the Catalog of World Fishes is difficult to use and is not recommended as a modern list of living sharks, especially to fisheries workers and others unfamiliar with shark systematics. Oftheapproximately350speciesoflivingsharksthatarecurrentlyknownabout48%ofthemaretomy knowledge of nousetofisheries; 25% areof limiteduse,20% are of considerable importance,and 7% aremajor fisheriesspecies.The'useless'categoryisprobablyoverlarge,dueinparttomylackoffisheriesinformationfor manyofthespecies.MyexperienceinanalyzingtheresultsofafisheriessurveyconductedbyFAOpriorto publishingtheFAOSpeciesdentificationSheetsforFishingArea34(seeCompagno,1981aforasummaryofthis survey as applied to sharks) convinced me that little-known species of sharks, especially deep-water species, are beingtakeninfisheriesatleastasabycatchand attheminimumarebeingprocessedforfishmealandoil. Formerlymanydeepwatersharkswereprimarilytakenbyscientificandexploratoryfisheriesexpeditionsand small-scaledeepwaterartisanalfisheries(suchasoffMadeiraandJapan),but,withtheadventofoffshore internationalfleetswithgiganticfactoryshipsandtrawlersaswellasspecializeddeepwatersharkfisheries aimingatprocessingsharkliveroilforsqualene,deepwaterspeciesarenowbeingincreasinglyfished.The48% 'useless'categoryincludesmanydeepwaterbenthicmembersofthefamilies Squalidae andScyliorhinidae,atleast someofwhicharequitelikelybeingtaken.twouldbemisleadingandinappropriatetolimitthepresentcatalogue to the known 'useful' species, because most if not all species of sharks are of potential use for fisheries (ifonlyforfishmealandoil);andavailablefisheriesdataissufficientlysketchytomakemestronglysuspectthat boththereporteddiversityofsharkspeciesbeingtakenandthereportedweightsofworldsharkcatchesarelower thanwhatisactuallybeingcaught.Quiteoftennational,regionalandworldsharkcatchstatisticsarenot segregatedbyspeciesandareoftenlumpedwithcatchesofbatoids.Oftenonlymajorfisheriesspeciessuchas pikeddogfish(Squalusacanthias)orporbeagle(Lamnanasus)arelistedseparatelyincatchstatistics(seeFAO Yearbook of Fisheries Statistics, 1983). - 1 -click for previous page feel also that a comprehensive catalogue of world sharks can serve as an invaluable educational tool to the fisheriesworkers,tosharkresearchesnotspecializingintaxonomy,topeopleencounteringsharksinthefieldand uncertainoftheirdangerorutility,andtotheinterestedpublicatlargeasaguidetothetaxonomicliterature andtothenumerousnamechangesandadditionsofspeciessincethelastcomprehensiveworksonsharkswere published.Therearenumeroustaxonomicsynonymsanddubiousspeciesnamesforsharksintheliterature, averaging.about1.7pervalidspeciesorabout5forevery3validspeciesdescribed,withasmanyas21foras inglespecies:theCarcharhinidae,withabout49validspecies,hasover150synonyms,.dubiousnamesandnomina nudaforspecies. ThepresentCatalogue listsall ofthesynonymsforsharkspecies, generaandfamilies,knownto me;andforspecieslistscommoncombinationsofgenericandspecificnamesthathaverecentlybeenusedbut are not considered valid. 1.1 PIan of the CataIogue This Catalogue is based on original work on various groups of sharks as well as my interpretation of data in theliterature.Someofthearrangementsoffamilies,generaandspeciesusedheredisagreewiththoseof previousworkers,butinsuchcasesthedisagreementsarediscussedorreferenceismadetodiscussionsinthe literature.Thesystematicarrangementusedhereisbasedonmyearlierworks(Compagno,1973c,1977,1979), whichdividesthesharksintoeightmajorgroupsororders.Therelationshipsofthesharkorderstoeachotherand totheraysorbatoids(Batoidea)issomewhatcontroversialatpresent,butinlieuoffurtherworkthelinear arrangement of orders and families in Compagno (1973c) is largely adhered to in this Catalogue (Fig. 1). Ordersarethehighesttaxonomicgroupsdealtwithhere,andmanyoftheirsynonymsarelistedeventhough thepresentnternationalCodeofZoologicalNomenclaturedoesnottreatgroupshigherthanthefamily-group level(superfamiliesandbelow).ThenomenclatureforordersisthatofCompagno(1973c),witholderandnewer equivalentslistedfromoldesttonewest.Theordersaresuffixedwith-iformesfollowingcommonichthyological practiceatpresent.Familiesaresuffixedwith-idae,theuniversalendingforzoologicalfamilies.Otherlevels betweenorders,families,generaandspeciesarenotcoveredhere.Subgeneraarediscussedundertheir appropriategenerabutspeciesarenotgroupedundersubgeneraandgivenparentheticalsubgenericnamessuchas Somniosus(Rhinoscymnus)rostratus,evenwheresubgeneraareconsideredvalid, so asnotto eliminatetheutility oflistingspeciesalphabeticallywithingenera.Subspeciesarelistedinthesynonymiesoftheirspeciesbutare not given separate cover. Families,generaandspeciesareprovidedwithcitationsfortheirauthor,yearofpublication,referenceand pagination(illustrationsalsoincludedforspecies),whilesynonymsandcombinationsofgeneraandspecies recentlyusedbutatvariancewiththepresenttextarecitedwithauthoranddateonly.Thepapersindicatedin thecitationsoffamilies,generaandspeciesaswellasthoseinwhichsynonymsandcombinationshaveappeared arenotnecessarilylistedinthebibliography,butliteraturecitedinotherpartsofthetextisincludedthere.The bibliographycoversawideselectionofreferencesusedinwritingthecatalogue,butisnotintendedtobeall-inclusive. Theinformationpertainingtoeachfamily,genusandspeciesisarrangedroughlyintheformusedina previouscatalogueinthisseries(Holthuis,1980),butwithconsiderablemodifications.Thefamilyaccounts includethevalidmodernformofthefamilynamewithauthorandyear;theoriginalcitationofthefamilyname withitsauthor,year,referenceandpagination;theFAOfamilyvernacularname;familysynonymswithname, authorandyear;afieldmarksanddiagnosticfeaturesofmembersofthefamily;anaccountofthenatural historyofthefamilyunderHabitat,DistributionandBiology;asynopsisoftheutilityofmembersofthefamilyto fisheries, under nterest to Fisheries; and a Remarks section. GenericAccountsincludethevalidmodernformofthegenusnamewithauthorandyear;theoriginal citation of the genus or subgenus, with its author, year, reference and pagination, and, if a subgenus, the original genus namewithauthorand yearthatthe subgenuswas placed in;the type speciesandmeansofdesignating it (forexample,byoriginaldesignation,monotypy,absolutetautonymy,orsubsequentdesignation);generic synonyms, with their rank (genus, subgenus, or other genus-group ranking such as W.H. Leigh-Sharpe's 'pseudo-genus'),author,year,andgenustheyweredescribedinifsubgeneraorequivalents;sometimesfieldmarksif genera are large; diagnostic features of the genus; a key to species; and a Remarks section. SpeciesAccountsincludethevalidmodernnameofthespecies,withauthoranddate;theoriginalcitation ofthespecies(orsubspecies),withitsauthor,year,referencepagination,andillustrations;alateralview illustration,andoftenotherusefulpartsofthesharkinquestion;theEnglish,FrenchandSpanishFAOspecies vernacularnames;theholotype,syntypes,lectotypeorneotypeofeachspecies(paratypesarenotlistedinthe present account), including the total length and sex of the specimen, its institutional deposition, and its catalogue number;thetypelocality(inthepresentcase,thelocationanddepththattheholotype,syntypes,lectotypeor neotypewerecaught);speciessynonyms,includingtheirnames,authorsanddates;asectionlistingother scientificnamesrecentlyinuse;fieldmarks;diagnosticfeatures(exceptinmonotypicgenera);geographical distribution,includingamap;habitatandbiology;size;interesttofisheries;sometimeslocalnames;literature; and a remarks section. - 2 -SHARKSno anal finanal fin presentbody flattened, raylike, mouth terminalbody notraylike,mouth ventral6 or 7 gill slits, one dorsal fin5 gill slits,2 dorsal finssnout short, not sawlikeno fin spinedorsal fin spinesmouth behindfront of eyesmouth well in front of eyesFig. 1 Higher Classification of sharks ( Orders )SQUATINIFORMESnictitating eyelids, spiral orscroll intestinal valve~~!".no nictitating eyelids,ring intestinal valveSOUALIFORMESHEXANCHIFORMESHETERODONTIFORMESCARCHARHINIFORMESPRISTIOPHORIFORMESLAMNIFORMESORECTOLOBIFORMESsnout elongated, sawlike- 3 - Synonymy:includesonlytruetaxonomicsynonymsofthevalidfamily,genusandspeciesgiven.Forspecies,anothercategory,OtherScientificNamesRecentlyinUse,isprovidedforcommonmisidentificationsofagiven specieswithanothervalidspeciesforexample,CarcharhinusbrachyuruswasuntilrecentlyoftentermedC. remotus,butthelatterisajuniorsynonymofC.acronotusaswellascommonlyusedcombinationsthatplacea

valid species in different genera (for example, Odontaspis taurus for Eugomphodus taurus). FAOFamiIyandSpeciesVernacuIarNames:English,FrenchandSpanishnamesforeachfamilyandspecies,primarilyforusewithinFAO,wereselectedbythefollowingcriteria:(a)eachnameappliestoasingle familyorspeciesworldwide;(b)thenameconformswithFAOspellingnomenclature;(c)thenameconformsto priorusagewhenpossible.FAOnamesarenotintendedtoreplacelocalspeciesnames,butarenecessaryto overcome the confusion caused by the use of a single name for more than one species or several names for one species.ThevernacularnamesusedhereconformwithpriorFAOusageandwhenpossibleandappropriate nationalandinternationalchecklistsandreviewsofspeciessuchasWhitley(1940),Shiino(1972,1976),Hureau& Monod(1973),Smith(1975),Robinsetal.(1980)andLindberg,Heard&Rass(1980).TheFrenchnameswere selectedjointlywithDrJ.-C.Du-ro,nstitutScientifiqueetTechniquedesPchesMaritimes,Ministredela Marine Marchande, La Rochelle, France. The names selected correspond to official French species nomenclature currentlybeingestablishedbytheDirectiondesPchesMaritimes.TheselectionofSpanishnamespresented considerabledifficultiesduetothelackofdenominationsformanyspecies.Whereverpossible,the"official" Spanish names adopted by F. Lozano in his book "Nomenclature ictiologica", Madrid 1963, were used. Theterm'shark'isvirtuallyuniversallyusedasacatchallterminEnglishforallmembersoftheClass Chondrichthyesthatarenotbatoidsorchimaeras.TheFrench'requin'andSpanish'tiburon'arecomparable generalterms.Severalnamesnotincorporating'shark'oritsFrenchorSpanishequivalentsareusedforsharks and are not used for other fishes; these include the English 'dogfish' (a regional name for the bowfin, Amia calva, 'freshwaterdogfish'),'smoothhound','tope','porbeagle',and'hammerhead'('wobbegong'isadaptedfroman AustralianAboriginaltermforsharksofthegeneraEucrossorhinusandOrectolobus).French'roussette', 'emissole','renard','milandre','marteau',and'griset',andSpanish'gato','cazon','tollo','pintarroja',and'cornuda', are similar terms for certain kinds of sharks. Usageofgeneralvernacularnamesfordifferentkindsofsharksvariesfromcountrytocountry.'Catshark' isusedformembersoftheScyliorhinidaeandsometimesrelatedfamilies(suchasProscylliidae)intheUnited States,butalsoforvariousorectoloboidsinAustralia.'Dogfish'isvariablyusedformembersofthefamilies Squalidae('spinydogfishes'),Scyliorhinidae(especiallyScyliorhinus),andTriakidae('smoothdogfishes',Mustelus spp.).'Sandshark'mayrefertoOdontaspididae(especiallyEugamphodustaurus,the'sandtigershark'ofthe easternUSA,called'ragged-toothshark'inSouthAfricaand'graynurseshark'inAustralia),toTriakidae (Mustelus spp.), or even to the batoid Rhinobatidae (guitarfishes). n the present catalogue 'catshark' is restricted tomembersoftheScyliorhinidaeandProscylliidae('falsecatsharks'aremembersofthePseudotriakidae), 'dogfish' to the Squalidae, and 'sand sharks' in the form of 'sand tiger shark' to the Odontaspididae. Orectoloboid 'catsharks' aretermed'carpetsharks',and'sandsharks'and'smoothdogfishes'ofthetriakidgenus Mustelusare termed 'smoothhounds' (except for M. antarcticus, the Australian 'gummy shark'). Keys, FieId Marks, and Diagnostic Features : Thesesectionsincludeidentificationdataindifferent forms. Keys to orders, families, genera and species are standard dichotomous biological keys that are followed in stepsofalternatechoicestosingleoutthetaxacovered.DiagnosticFeaturesarecomprehensivelistsof charactersattheordinal,familial,genericandspecieslevel,withthecharacterchoicegenerallylimitedto externalcharactersbecauseofspaceconsiderationsandtheirprimarypurposeofidentificationratherthan indicationofrelationships.TheDiagnosticFeaturessectionsarehierarchical,withcharactersattheordinallevel notduplicatedatthefamily,genusandspecieslevel.Monotypicorderswithonefamily(suchasPristiophori-formes),monotypicfamilieswithonegenus(Chlamydoselachidae)ormonotypicgenerawithonespecies (Carcharodon) all have the Diagnostic Features section present only in the highest taxon covered. n a monotypic order,DiagnosticFeaturesareomittedintheaccountofitssinglefamily;inamonotypicfamily,theyareomitted from its single genus; and in a monotypic genus, they are omitted from its single species. FieldMarksincludeafewobviouscharactersofuseinfieldidentification,extractedfromDiagnostic Featuresatvariouslevels,butincludedinaseparatesection.FieldMarksarelistedattheordinal,familialand species level, and occasionally the generic level in cases of large genera with many species. The arrangement of FieldMarkcharactersissemihierarchicalandmayincludecharactersfromahigherlevelsuchasanorderin lower level taxonomic accounts such as those of species, depending on their utility. AnexampleofthedifferentapplicationofDiagnosticFeaturesandFieldMarksisindicatedwiththesevengillshark, Heptranchiasperlo.StartingwiththeOrder Hexanchiformes, Diagnostic Features applicableto it are given at decreasing hierachical levels through the Family Hexanchidae and Genus Heptranchias (a monotypic genus).However,thespeciesaccountofH.perloalsohasashortFieldMarkssection,"Anarrow-headed,big-eyed,smallseven-gilledsharkwithonedorsalfin",thatcansufficetoidentifyitwithoutadditionalinformation, althoughthisisavailableintheDiagnosticFeaturessectionsasneeded.nsomelargefamilieslikethe CarcharhinidaetheFieldMarksforaneasilyrecognizedspecieslikeCarcharhinuslongimanusmaynotrepeat familial and ordinal characters but merely indicates its family and unique characters. - 4 -GeographicaIDistributionandMaps:Sharksareprimarilymarineorganisms,butanumberofspeciesreadilyenterbrackishtoalmostfreshwaterestuaries,lagoonsandbays;afewspeciesofthefamily Carcharhinidaeoccurfarupriversandinfreshwaterlakeswithconnectionstothesea.Distributionsfornearly allspeciesaregivenbylistingthecountriesoffthecoastsofwhichthesharksoccur.nthecaseoftwospeciesofcarcharhinids(CarcharhinusleucasandGlyphisgangeticus)thatareknownfromverifiablerecordsfrom entirelyfreshwaterpartsofriversandfreshwaterlakes,thenamesofmajorriversystemsandlakeswherethey occurarenoted.TherearevariousfreshwaterrecordsofothermembersofthefamilyCarcharhinidaeandeven someotherfamilies(includingthezebrasharkfamily,Stegostomatidae)butmanyoftheserecordsmaybefrom almostfreshwaterlowerreachesofriversandestuariesandmayindicatethespeciesinvolvediseuryhalinebut incapableofexistinginfreshwaterfarfromtheseaascanC.leucas,apparentlyG.gangeticus,andamong batoidssomesawfish(Pristidae),potamotrygonidstingrays,andsomedasyatidstingrays.Someofthese carcharhinidfreshwaterrecordsmaybebasedonC.leucasorGlyphisspeciesratherthanthespeciesindicated (such as C. melanopterus or C. hemiodon). ncompilingdistributionaldataandpreparingmaps,itwasnotedthatthedistributionsofmanywide-ranging coastalspeciesareveryspottyasknownatpresent,especiallywithspeciesoccurringinthendianOceanand westernPacific.nmanycasesgapsindistributionmaynotindicateabsenceofagivenspeciesbutabsenceof knowledge.Continentalslopesharkfaunasarepoorlyknownformuchoftheworld,andanumberofdeepwater speciesmayhavewiderrangesthanarecurrentlyknown.Thelocalitydataintheliteratureandonspecimen labels is often very general and imprecise; and even with detailed oceanographic data trawl hauls may be of such longdurationthatlocationsareapproximate.Hencedistributionaldataandmapspresentedhereistobe consideredasroughapproximationsofdistribution.Mucheffortwasmadetoscreenoutmisinformationanshark distribution, based on misidentifications of species, to the cost of presenting distributional lists and maps that are spotty if more accurate. An extreme example is discussed in detail under Glyphis gangeticus (Carcharhinidae). HabitatandBioIogy:Thissectioncoversinformationonnaturalhistory.Theknowndepthrangeofthespecies(inmeters),positioninthewatercolumn,typeofsubstrateoccupied,andpreferencesrelativetocoasts are noted when available. n most species data on salinity, oxygen content, and specific temperature of the water in which they occur is not available or not in an easily usable form and has not been compiled here. Also included aredataonpopulationstructureanddynamics,reproduction,behaviour,ageandgrowth,feedinganddangerto people.ncompilingthedataitwasnotedthatveryfewspeciesarebiologicallywellknown,andeveninthepikeddogfish(Squalusacanthias),thebest-knownoflivingsharks,thereareareasofitsbiologythatarevery poorly known (such as its ethology). There is a bias in available natural history data towards reproductive biology, feeding, shark attack and fisheries-related subjects, and correspondingly little on ecology and behaviour. Size:Allsizedataisgivenastotallengths;thisisthemeasurementmostoftenusedasanindependentvariableandstandardmeasurementinthesharkliterature,althoughparticularlyinfisheriespapersprecaudal lengths,forklengths,andothermeasurementshavebeenusedfromchoiceornecessity.Unfortunatelyshark workershavenotagreedonastandardmethodofmeasuringtotallength,sototallengthsfromdifferentsources intheliteraturemaynotbestrictlycomparable.preferandadvocateasastandardmethodadirect measurement,inwhichthesharkisheldbellydownwithitsdorsalcaudallobedepressedintolinewithitsbody axisandtotallengthmeasuredasapointtopointdistance(notoverthecurveofthebody)fromthesnouttipto the tip of the dorsal caudal lobe (see also Compagno, 1979, 1981a). This method lends itself readily to quick use ofafishboardwithaperpendicularfrontbarorplatetoindexthesharks'ssnoutagainst,ametreortwometre ruleroffoldingruleslippedundertheshark,orevenasteelorclothtape,acidavoidsthetroubleofcomputation and possible errors and loss of data. Acomparablecomputationalmethodaddingthelengthsfromsnouttiptocaudaloriginandofthedorsal caudalmarginisadvocatedbySadowsky(1968).Bigelow&Schroeder(1948)andV.Springer(1964)measuredtotal lengthfromthesnouttipalongthebodyaxistoaverticalprojectionfromthetipofthedorsalcaudallobewith thecaudalfinina'naturalposition'.Bass(1973)advocatedacomputationalmethodwhichadds,thelengthfrom snouttiptocaudalorigintoanumbercomputedbymultiplyingthelengthofthedorsalcaudalmarginbya constant (1.0 or less; 0.97 and 0.80 were the numbers used) that corrects for the 'natural' angle of the caudal axis tothebodyaxisindifferentspecies.Themethodadvocatedheredispenseswithallcomputationfordetermining total length and avoids arbitrary constants to correct for supposed 'natural positions' of the caudal axis as well as thedifficultiesinobtainingaccurateverticalprojectionsfromarbitrary'naturalpositions'.Also,withthepresent methodacomparablemeasurementcanbeobtainedforallsharks,raysandchimaeras;incontrast'natural position' methods arbitrarily generate incompatable total lengths for different sharks. Totallengthdatapresentedincludesmaximumsize,sizeatmaturity(insomecases,asizerangeat maturity,whenabundantdatawasavailable)andmaximumsizeforbothsexes(assexualdimorphisminsizeis nearlyuniversalamongsharks,withfemalesusuallyattaininglargersizesthanmales),andsizeatbirthor hatching.Sometimessizewheneitherorbothsexesmatureisnotknown,inwhichcasesreportedminimumand maximum sizes of adult individuals are given. n some cases maximum size exceeds that recorded for either sex, in which case the sex of the outsized individual or individuals representing the maximum size measurements was notrecorded.nsomepoorlyknownspeciesonlyimmatureindividualsareknown,inwhichcasethehypothetical maximumsizeisalmostcertainlylargerthantheknownimmaturemaximum(nocasesofadultsharksthatare considerably smaller than large immatures are known, unlike some other vertebrates) - 5 - nasmallnumberofspecies length-weightequationsarepresented,usuallyoftheformWT=a+ TLexpb, where Wt is weight, a and exp b are constants, and TL is total length. InteresttoFisheries:Asnotedabove,fisheriesdataforsharksisoftensketchyandcombinedforanumber of species. n this section data on localities of fisheries, gear used, and uses of the particular species are notedwhenavailable.Catchstatisticsonsinglespecieswereavailableforonlyafewsharksbutarenotedwhen available. LocaI Names :Localvernacularnamesareusuallynotlistedinthepresentversionofthiscatalogue because these names are spottily covered in the literature; no worldwide survey of current usage exists, although Fowler(1966-1969), Shiino (1976)and Lindberg, Heard & Rass(1980)have compiledvernacularnamesfromthe literatureforanumberofspecies.Duetotimeconstraintsasurveyorregionalvernacularnamescouldnotbe attemptedandcompiledhere,butafewalternateregionalnames(mostlyinEnglish)arelistedwhenthesehave sufficientlywideandconsistantusagesothattheirinclusionisveryhelpful.Examplesincludeschoolshark (Australia),tiburonvitaminicoorvitaminshark(UruguayandArgentina)andsoupfinshark(PacificUSAand Canada)forthetopeshark(Galeorhinusgaleus);andragged-toothshark(SouthAfrica)andgraynurseshark (Australia) for the sand tiger shark Eugomphodus taurus). Literature : Referencescitedhereincludesspecificworkswithimportantinformationaswellas comprehensive accounts on species, and is not intended as an extensive bibliography. Remarks:mportantinformation,especiallyonsystematicsandnomenclature,notrepresentedelsewhereare given here. AcknowIedgements Thiscataloguehasbeenoflonggestation,stemmingfromunpublishedchecklistsofsharks,raysand chimaerasthewriterhasmaintainedoverthepasttwoandahalfdecades.Manypeoplehavehelpedmeduring thecourseofthiswork.wouldespeciallyliketothankW..Follett,oftheCaliforniaAcademyofSciences,San Francisco,California,forencouragementonmyinitialchecklistsmany,manyyearsago.Especialthanksgoto three of my colleagues in elasmobranch research, Shelton P. Applegate (Los Angeles County Museumandnstituto deGeologia,CiudadUniversitaria,MexicoCity),J.A.F.Garrick(DepartmentofZoology,VictoriaUniversity, Wellington,NewZealand),StewartSpringer(NationalMarineFisheriesService,USA,andMoteMarine Laboratory,Sarasota,Florida),allofwhomhavebeenextremelyhelpfulandinspirationalovertheyears.Thanks alsotoDrWarrenC.FreihoferoftheDivisionofSystematicBiology,StanfordUniversity,CaliforniaAcademyof Sciences and Tiburon Center for Environmental Studies, for much support and encouragement. ThepresentcatalogueowesanenormousamounttotheuntiringandtitaniceffortsofDrW.Fischer, Fisheries Resources and Environment Division, FAO Fisheries Department, who inspired me to make it a far more usefulworkthanwouldhavebeenmyinclinationasaprofessionaltaxonomist;andcausedmetolearnan enormousamountintheprocessofwritingit.SpecialthanksgotoOrBernardJ.Zahuranec,oftheUSOfficeof NavalResearch,OceanicBiologyProgram,forprovidingsupportforthewritingofthecatalogueoverthepast threeyearsintheformofaresearchcontracttothewriterattheTiburonCenterforEnvironmentalStudies,San FranciscoStateUniversity,Tiburon,California,andforarrangingfundingforacircumglobalresearchtripin 1982.Prof.MargaretM.SmithandDrMichaelBrutonarrangedforaseniorresearchfellowshipforthewriterat theJ.L.B.SmithnstituteofchthyologyforamongotherprojectsthecompletionofthisCatalogue.Thanksalso toDrSamuelE.GruberandtheAmericanElasmobranchSocietyforsupportforthepublicationofthecatalogueintheUnitedStates.DrCorneliaE.NauenoftheFisheriesResourcesandEnvironmentDivision,FAOFisheries Department,wasextremelyhelpfulintheorganizationofthefinalversionofthecatalogue,whichwentthrough severalchangesinformatduringthepastfiveyears.ThanksalsotothestaffofDrW.Fischer'sprogrammefor identificationoffisheriesspeciesatFAOfortheirefforts,particularlyforthefineartisticworkofEmanuela D'Antoni,Pierluigisola,PaoloLastrico,andOlivieroLidonniciintranslatingoftendifficultmaterialfromthe literatureaswellasthewriter'sresearchdrawingsandmapsintotheillustrationsinthiscatalogue;andtoMrs GiuliaSciarappa-Demurofortypingthefinalmanuscript.MrsPaulaSmith,MsBarbaraNabors,andMsDale WhiteoftheTiburonCenterforEnviromentalStudieshelpedinthetypingandcopyingofversionsofthe manuscript and the organization of the bibliography. - 6 -Specialthanksalsotothefollowing:GeorgeS.Myers,NormanK.Wessells,PaulR.Ehrlich,Donald Kennedy,RobertT.Schimke,JohnH.Thomas,andMargaretA.SharpoforformerlyoftheDepartmentof BiologicalSciences,StanfordUniversity,Stanford,California,wherethewritercompletedadoctoraldegreeand learnedmuchofuseinthiscatalogue;SusumuKatoandSusanSmith(NationalMarineFisheriesService,Fisheries Laboratory, Tiburon, California); Bruce Welton (University of California, Berkeley, Los Angeles County Museum of NaturalHistory,andChevronOilFieldResearchCo.,LaHabra,California);RobertLea(California DepartmentofFishandGame);GregorCaillietandDaveEbert(MossLandingMarineLaboratories);Bobb Schaeffer,DonnRosenandGarethNelson(AmericanMuseumofNaturalHistory,NewYork);JohnR.Paxton (Australian Museum, Sydney); Alwynne Wheeler, Peter Whitehead, Oliver Crimmen, and David Ward of the British Museum(NaturalHistory),London;W..Follett,WilliamD.Eschmeyer,TysonR.Roberts,LillianJ.Dempster, RobertP.Dempster,Tomiowamoto,andPearlSonoda(CaliforniaAcademyofSciences);E.G.Silas,K.K. Appukuttan,andM.E.Rajapandian(CentralMarineFisheriesResearchnstitute,Cochin,ndia);L.W.Filewood (DepartmentofAgriculture,StockandFisheries,Konedobu,PapuaNewGuinea);RainerZangerl(Departmentof Geology,FieldMuseumofNaturalHistory,Chicago);MargaretSmith,MichaelBruton,andPhillipC.Heemstra (J.L.B.Smithnstituteofchthyology,Grahamstown,SouthAfrica);RobertJ.LavenbergandCammSwift(Los AngelesCountyMuseumofNaturalHistory);MyvanwyM.Dick(MuseumofComparativeZoology,Harvard University,Cambridge,Massachusetts); LouisGaribaldiand Don Reed (Marine World,Redwood City,California); John Moreland (National Museum Welligton, New Zealand); John Bass and John Wallace (Oceanographic Research nstitute,.Durban,SouthAfrica);DarioJ.Guitart(nstitutodeOceanologia,AcademiadeCienciasdeCuba, Havana);thelateJ.L.B.Smith(RhodesUniversity,Grahamstown,SouthAfrica);C.G.Alexander,Robert. Bowman,MargaretG.Bradbury,andMichaelJocelyn(SanFranciscoStateUniversityandTiburonCenterfor EnvironmentalStudies);thelateCarlL.HubbsandRichardH.Rosenblatt(ScrippsnstitutionofOceanography,La Jolla,California);LeslisW.Knapp(SmithsonianOceanographicSortingCenter,Washington,D.C.);JohnD. McEachran(TexasA&MUniversity);C.RichardRobins(nstituteofMarineSciences,UniversityofMiami); ReeveM.Bailey(UniversityofMichiganMuseumofZoology,AnnArbor);StanleyH.WeitzmanandVictorG. Springer(DivisionofFishes,USNationalMuseumofNaturalHistory);BruceB.Collette(NationalMarineFisheries Service,SystematicsLaboratory,USNationalMuseumofNaturalHistory,Washington,D.C.);VictorSadowsky (nstituto Oceanografico, Universidade de Sao Paulo, Sao Paulo, Brazil); Toru Taniuchi (Department of Fisheries, UniversityofTokyo);Wolf-ErnstReif(nstitutandMuseumfrGeologieandPalontologiederUniversitt Tbingen,WestGermany);GeraldAllenandJ.BarryHutchins(WesternAustralianMuseum,Perth,Australia); LeightonR.Taylor,Jr(WaikikiAquarium,Honolulu,Hawaii);JohnC.Cordell(WoodsHoleOceanographic nstitution, Massachusetts); P.K. Talwar (Zoological Survey of ndia, Calcutta). John W. Shipman of Socorro, New Mexicogavemuchadviceoncomputersandtext-editingusefulforthiswork,andpreparedacomputerizedversion oftheinitialchecklistmanuscript.ThanksalsotoTomCrowellofComputerlandofMarin,California,foradvice andsupportinobtainingandmaintainingERS,thewriter'saugmentedCOMPAQpersonalcomputeronwhich muchrevisionaryworkonthecataloguewaswrittenwiththeWordStartext-editinqprogramme.ammostthankful for the help offered by all of these people and by others not mentioned, without whom this account could not have been written. - 7 -1.2Technical Terms and Measurements1.2.1 Picture Guide to Terminology of Sharkslabialfurrowseyespectoral finsubocular pocketfirst dorsal findorsal finpreventral marginventral lobecaudal finventral tipcaudal finterminal marginsubterminal margin.subterminalnotchupper postventral marginposterior notchlower postventral margin- 8 -snoutnostrileyespiraclespinedorsal finsecond dorsal finprecaudal pitcaudal keelanal finclasper (males)pelvic finlateral viewupper origingill openingsmouthdorsal margindorsal lobeterminal lobeposterior tiplower originupper eyelidnotchnictitatinglower eyelidsecondarylower eyelidspineapexposteriormarginfree rear tipinner margininsertionbaseoriginanteriormarginanterior nasal flap liftednasoral groovemouthsymphyseal grooveanterior nasal flaplower labial furrow pectoral finexcurrent aperturenostrillabial furrowmouth corner - 9 -snoutnostriltrunkgill openingheadprecaudaltailanal fincaudal finpreanal ridgesventpelvic fin(female, no claspers)ventral viewpectoral finmouthincurrent apertureanterior nasal flapexcurrent aperturediagrammatic orectoloboid headincurrentaperturecircumnarial groovecircumnarial foldbarbelupper labial furrowinsertioninner marginfree rear tipposterior marginapexanteriormarginoriginbaselabial foldposterior nasal flapPD2PRCFORTOTPALPRN PRENARIAL LENGTHPORPREORAL LENGTHEYL EYE LENGTHEYH EYE HEIGHTING INTERGILL LENGTHGS1FIRST GILL SLIT HEIGHTGS2 SECOND GILL SLIT HEIGHTGS3THIRD GILL SLIT HEIGHTGS4 FOURTH GILL SLIT HEIGHTGS5FIFTH GILL SLIT HEIGHTGS6SIXTH GILL SLIT HEIGHTGS7SEVENTH GILL SLIT HEIGHTPlAPECTORAL ANTERIOR MARGINPlRPECTORAL RADIAL LENGHTPlBPECTORAL BASEP1I PECTORAL INNER MARGINPIPPECTORAL POSTERIOR MARGINPIHPECTORAL HEIGHTSODSUBOCULAR POCKET DEPTH1.2.2 Measurements Used for SharksTOT TOTAL LENGTHPP2 PREPELVIC LENGTHFOR FORK LENGTHSVL SNOUT-VENT LENGTHPRC PRECAUDAL LENGTHPALPREANAL LENGTHPD2 PRE-SECOND DORSAL LENGTHIDS INTERDORSAL SPACEPD1 PRE-FIRST DORSAL LENGTHDCS DORSAL-CAUDAL SPACEHDL HEAD LENGTHPPS PECTORAL-PELVIC SPACEPG1 PREBRANCHIAL LENGTHPASPELVIC-ANAL SPACEPSP PRESPIRACULAR LENGTHACS ANAL-CAUDAL SPACEPOBPREORBITAL LENGTHPCAPELVIC-CAUDAL SPACEPP1 PREPECTORAL LENGTHVCL VENT-CAUDAL LENGTH- 10 -======================================PD1HDLPGIPSPPOBPP1PP2PPSSVL VCLEYLEYGP1BINGP1IP1LP1RP1ASODPRNPORDCSIDSGS5GS1P1HP1Pcross-sectionthrough eyePCAACSPAS- 11-CDM = DORSAL CAUDAL MARGINCPV = PREVENTRAL CAUDAL MARGINCPU = UPPER POSTVENTRAL CAUDAL MARGINCPL = LOWER POSTVENTRAL CAUDAL MARGINCFW = CAUDAL FORK WIDTHCFL = CAUDAL FORK LENGTHCST = SUBTERMINAL CAUDAL MARGINCSW =SUBTERMINALCAUDAL MARGINCTR = TERMINAL CAUDAL MARGINCTL = TERMINAL CAUDAL LOBEDIL = FIRST DORSAL LENGTHD1A = FIRST DORSAL ANTERIOR MARGINDIB = FIRST DORSAL BASED1H = FIRST DORSAL HEIGHTD1I = FIRST DORSAL INNER MARGINDIP = FIRST DORSAL POSTERIOR MARGIND2L = SECOND DORSAL LENGTHD2A = SECOND DORSAL ANTERIOR MARGIND2B = SECOND DORSAL BASED2H = SECOND DORSAL HEIGHTD2I = SECOND DORSAL INNER MARGIND2P = SECOND DORSAL POSTERIOR MARGINP2L = PELVIC LENGTHPZA = PELVIC ANTERIOR MARGINP2B = PELVIC BASEP2H = PELVIC HEIGHTP2I = PELVIC INNER MARGIN LENGTHP2P = PELVIC POSTERIOR MARGIN LENGTHHDH = HEAD HEIGHTTRH = TRUNK HEIGHTABH = ABDOMEN HEIGHTTAH = TAIL HEIGHTCPH = CAUDAL PEDUNCLE HEIGHTcaudal finANL = ANAL LENGTHANA = ANAL ANTERIOR MARGINANB = ANAL BASEANH = ANAL HEIGHTANI = ANAL INNER MARGINANP = ANAL POSTERIOR MARGINDPO = FIRST DORSAL MIDPOINT-PELVIC ORIGINPDI = PELVIC MIDPOINT-FIRST DORSAL INSERTIONPDO = PELVIC MIDPOINT-SECOND DORSAL ORIGINDAO = SECOND DORSAL ORIGIN-ANAL ORIGINDAI = SECOND DORSAL INSERTION-ANAL INSERTIONPDICPLCPVCSWHDHD2HCTLCPUCTRCFWCFLCSTCDMD1AD1HD1PD1BD1LD1ID2LD2BD2ID2AD2PP2LP2BP2IANLANBANIP2HANAP2AANPP2PANHtrunkPDODPI DPODAO DAITRHABHTAHCPHMOL = MOUTH LENGTHMOW = MOUTH WIDTHULA = UPPER LABIAL FURROW LENGTHLLA = LOWER LABIAL FURROW LENGTHNOW = NOSTRIL WIDTHINW = INTERNARIAL SPACEANF = ANTERIOR NASAL FLAP LENGTHESLCLO = CLASPER OUTER LENGTHCLI = CLASPER INNER LENGTHCLB = CLASPER BASE WIDTHCLIclasperINO = INTERORBITAL SPACESPL = SPIRACLE LENGTHESL = EYE SPIRACLE SPACEHDW = HEAD WIDTHTRW = TRUNK WIDTHABW = ABDOMEN WIDTHTAW = TAIL WIDTHCPW = CAUDAL PEDUNCLE WIDTH- 12 -INWANFMOWLLAULANOWGIRMOLnostrilangle of mouthventral viewGIR = GIRTHINOSPLdorsal viewHDWTRWABWTAWCPWCLOclick for next page - 13 - 2 ORDER HEXANCHIFORMES - FRILLED AND COW SHARKS Order Hexonchiformes Compagno, 1973c, J.Linn.Soc.(Zool.), Lond., 53, suppl. 1:37 p. Synonymy :Superorder Carcharhini, Order Hexanchida, Suborder Hexanchoidei:Glikman,1967(inpart). SuperorderChlamydoselachii,OrderChlamydoselachida:Glikman,1967(inpart).OrderChlamydoselachiformes: Rass&Lindberg,1971(inpart),Applegate,1974(inpart).SuborderChlamydoselachoidea:Bigelow&Schroeder, 1948(inpart).OrderDiplospondyli:Jordan&Evermann,1896,Fowler,1941,Smith,1949.SuborderGalei:Gill, 1872(inpart).OrderHexanchea:White,1936,1937,Whitley,1940(inpart).OrderHexanchida:Fowler,1967a. SuborderHexanchida:Matsubara,1955.OrderHexanchiformes:Berg,1940,Berg&Svedovidov,1955,Arambourg &Bertin,1958,Patterson,1967,Lindberg,1971,Rass&Lindberg,1971(inpart),Applegate,1974(inpart), Nelson,1976,Chu&Wen,1979.SuborderHexanchiformes:Budker&Whitehead,1971.OrderHexanchaidea: Schultz&Stern,1948.SuborderHexanchoidea:Romer,1966."Group"Hexanchoidei:Garman,1913.Order Notidani:Jordan,1923(inpart).SuborderNotidani:Goodrich,1909.SuborderNotidaniformes:LozanoyRey, 1928,Bertin,1939.SuborderNotidanaidea:Romer,1945,Bigelow&Schroeder,1948(inpart),Norman,1966. "Di vi si on"Noti danoi dei :Regan,1906.SuborderNoti danoi dei :Engel hardt,1913.OrderOpi stharthri :Gi l l ,1893. (?Suborder)Palaeonotidani:Hasse,1879.OrderPternodonta:Gill,1884(inpart).OrderSelachii:Blot,1969(in part). Order Selachophichthyoidi: Garman, 1884 (in part), Jordan, 1923 (in part), Whitley, 1940 (in part). FieId Marks : 6 or 7 pairs of gill slits, one dorsal fin, anal fin. DiagnosticFeatures:Trunkcylindricalorsomewhatcompressed,notflattenedandraylike.Headconicalto sl i ghtl ydepressed,notexpandedl ateral l y;6or7pai rsofgi l l sl i tspresentonsi desofhead,wi ththe posteriormostinfrontofpectoralfinorigins;spiraclespresentandverysmall,wellbehindandabovelevelof eyes;nostrilswithoutbarbels,nasoralgroovesorcircumnarialgrooves,separatefrommouth,anteriornasalflaps shortandnotreachingmouth;eyesonsidesofhead,withoutnictitatinglowereyelids;snoutveryshortto moderatelylong,truncatedtoconical,notgreatlyelongatedorflattened,andwi thoutlateralteethorrostralbarbels;mouthlarge,archedandelongated,extendingwellbehindeyes;labialfurrowsreducedorabsent,on lowerjawwhenpresent;teethweaklytostronglydifferentiatedalongthejaws,withoutenlargedanterioror posteriorteethandwithoutagaporsmallintermediateteethbetweenanteriorandlateralteethintheupperjaw. Asinglespinelessdorsalfinpresent,withoriginoverorbehindpelvicfininsertions;pectoralfinssmallto moderatel ylarge,notexpandedandrayl ike,withouttriangularanteriorlobesthatcoverthegillslits;pelvicfi ns smalltomoderatelylarge,withventcontinuouswiththeirinnermargins,'analfinpresent;caudalfinwithalong dorsallobeandtheventrallobeshorttoabsent;vertebralaxiselevatedintothedorsalcaudallobe.ntestinal valve of spiral or ring type. Key to FamiIies 1a.Bodyelongatedandeel-like; mouth nearly terminal on head;3- cusped teeth in upper and lowerjaws (Figs 1,2) - Frilled sharks .......FamiIy ChIamydoseIachidae (one genus, Chlamydoselachus) upper and lower toothFig. 1 Chlamydoselachus 1b. Bodystouter,noteel-like;mouth subterminalonhead;high,long-cusped teeth in upper law, larger, comblike teeth in lower jaw (Figs3,4) - Cow sharks .................................. FamiIy Hexanchidae (see key to genera) ChlamydoselachusFig. 2 upper and lower toothFig. 3 Hexanchus Fig. 4 Hexanchusclick for previous page- 14 - 2.1 FAMILY CHLAMYDOSELACHIDAE Garman, 1884 CHLAM Chlamydoselachidae Garman, 1884, Bull.Essex nst., 16:8. Synonymy :None. FAO Names :En - Frilled sharks; Fr - Requins collerette; 5p - Tiburones anguila. Genera :A single genus, Chlamydoselachus Garman, 1884. Diagnostic Features : Body slender and eel-like, with prominent keels on abdomen.Headwith6pairsof gillslits,thelowerendsofthefirstgillslitsconnectedtoeachotheracrossthroat;snoutextremel yshort, truncated;mouthterminalonhead,veryl ong;teethali keinupperandlowerjaws,wi ththreestrongcuspsanda pairofintermediatecusplets,notsaworbladelike,posteriorteethnotminuteandgranular.Anal finlarge,larger than dorsal fin; caudal fin without a subterminal notch. ChIamydoseIachus Garman, 1884 CHLAM Chlam Genus : Chlamydoselachus Garman, 1884, Bull.Essex nst., 16:8. Type Species : Chlamydoselachus anguineus Garman, 1884, by monotypy. Synonymy :Genus Chlamydoselache Gnther, 1887 (emendation). ChIamydoseIachus anguineus Garman, 1884 CHLAM Chlam 1 ChlamydoselachusanguineusGarman,1884,Bull.Essexnst.,16:3,fig.Holotype:MuseumofComparative Zoology, Harvard University (MCZ) . Type Locality: Japan, probably southeastern Honshu. Synonymy : None. FAO Names : En - Frilled shark; Fr - Requin lzard; SP - Tiburn anguila. upper and lower toothunderside of head FieId Marks :Eel-likesharkwi th6gillslits,terminalmouthwithtricuspidteethinboth'jaws,andone dorsalfin. Diagnostic Features:See family. - 15 - GeographicaIDistribution:Wide-rangingbutspottily distributed.EasternAtlantic:FromnorthernNorway,the AtlanticSlopeoffnorthernScotlandandwesternreland, France,Spain,Portugal,Morocco,Mauritania,Madeira, Angola,northernNamibia,andpossiblytheeasternCapeof GoodHope,SouthAfrica.WesternndianOcean:OffSouth Africa.WesternPacific:OffJapan(southeasternHonshu), Australia(NewSouthWales),andNewZealand.Eastern Pacific. Southern. California, USA, northern Chile. Habitat and BioIogy :Benthicontheouterconti-nentalandinsularshelvesandupperslopes,atdepthsusually between120and1280m,butoccasionallytakenatthe surface. Arare,littleknowndeepwatershark.Ovoviviparous,numberofyoung8to12.Apparentl yreproducesallyearindeepwateroffJapan;gestationperiodprobablyverylong,onetotwoyears.Feedinghabitsunknown,buttheneedle-sharp,slender-cuspedteethofthissharksuggestfeedingondeepwatercephalopodsandbottomfishes. Not dangerous, but teeth can lacerate the hands of the unwary scientist examining its mouth. Size:Maximum196cm,sizeatbirthabout39cm;sizeatmaturityabout97cmformalesand135cmfor females. InteresttoFisheries:Ofl i ttl ei mportance,takeni ncidental l yi nbottomtrawl scatches,formerl ycaught with deepset longlines off Japan. Utilized for fishmeal and for human consumption. Literature :Gudger & Smith (1933);Gudger (1940);Bass,d'Aubrey&Kistnasamy(1975c);Cadenat& Blache (1981); Compagno (1981). 2.2 FAMILY HEXANCHIDAE Gray, 1851HEX Tri beHexanchi naGray,1851,Li stFi shBri ti shMus.,Part,Chondropterygi i ,Bri t.Mus.(Nat.Hi st.):67 (Family Squalidae). Synonymy :Subfamily Notidanini Bonaparte, 1838 (Family Squalidae);Family Hexeptranchidae Garman, 1913; Family Heptranchidae Barnard, 1925. FAO Names :En - Cow sharks, Sevengill sharks, Sixgill sharks;Fr-Requinssixetseptfentes branchiales, Requins griss, Requins perlon, Requins vaches; Sp - Caabotas.FieIdMarks:Cylindricalsharkswith6or7gillslits,subterminalmouthwithlarge,bladelike,comb-shaped teeth in lower jaw, and one dorsal fin. DiagnosticFeatures:Bodycylindricalandmoderatelyslendertostout,withoutkeelsonabdomen.Head with6or7pairsofgillslits,thelowerendsofthefirstgillslitsnotconnectedtoeachotheracrossthroat;snout shorttomoderatel yl ong,coni cal andsl i ghtl ypoi ntedtobroadl yrounded;mouthsubtermi nal onsnout, moderately long; teethwell-differentiatedinupperandlowerjaws,upperanterolateralssmall,narrow,witha maincuspandoftensmallercusplets,lowersverybroad,compressedandsawlike,withaseriesoflargecusplets andashorttoel ongatedcusp;posteri orteethsmal l andgranul aratcornersofdental bands.Anal fi nsmal l , smaller than dorsal fin; caudal fin with a strong subterminal notch. Habitat, Distribution and BioIogy : Cowsharkshaveaworldwidedistributioninborealandcold temperatetotropicalseas.Mostspeciesaredeepwaterinhabitantsoftheoutercontinentalshelves,upper continentalslopes,insularshelvesandslopes,andsubmarinecanyonsdowntoatleast1875mdepth,nearthe bottom or well above it; but also occurring in shallow bays, close inshore, and near the surface.Behaviour of thesesharksispoorlyknown:theyaresluggishtoactiveoftenstrong-swimmingsharksthatoccurnearthe bottom.Theyrangeinsizefromsmalltoverylarge,withvariousspeciesbetween1.4to4.8mmaximumtotal length at maturity. Theyfeedonrelativelylargemarineorganisms,includingothersharks,rays,awidevarietyofbonyfishes, crustaceans,andcarrion(includingmammalianmeat).Cowsharksareovoviviparous(aplacentalviviparous),and lack a yolk-sac placenta. - 16 - These sharks are apparently not implicated in unprovoked attacks an swimmers and divers, though they may bite aggressively during capture and can inflict severe lacerations when handled. One species (broadnose sevengill shark) has attacked divers in aquarium display tanks and two of the cowsharks (broadnose sevengill and bluntnose sixgillsharks)reachsufficientlylargesize(from2.9to4.7m)torankaspotentiallydangeroustopeopleinthe water. Interest to Fisheries : Cow sharks are relatively unimportant but regular components of shark fisheries and bycatchesofotherfisheriesintemperateandtropicalwaters,andaretakenbylinegear,bottomandpelagic trawls,andgillnets.Thesesharksareexcellentforhumanfoodandareutilizedfreshanddried-salted;theyarealso processed for fishmeal, oil, and leather. Some species are subject to sports fisheries in shallow waters. Literature:Garman(1913);Daniel(1928);Fouler(1941);Bigelow&Schroeder(1948);Garrick& Schultz (1963); Springer & Waller (1969); Garrick & Paul (1971); Bass, d'Aubrey & Kistnasamy (1975c). Remarks :The genus Heptranchias is morphologically divergent from other members of Hexanchidae and has sometimes been placed in its own family, Heptranchidae (Compagno, 1973c). Key to Genera and Species 1a.Sixpairsofgillopenings(Fig.1) (Hexanchus) 2a. Snout shorter, blunt and broad;lower jaw with 6 rows of large, Fig. 1combl i keanterol ateral teethoneachsi de;Dorsal fi nbasesepa-ratedfromuppercaudal fi nori gi nbyadistanceaboutequaltoor slightlygreaterthanitslength.Sizeverylarge,upto4.7mtotallength (Fig. 2) - Broadnose sixgillshark ............................................... Hexanchus griseus Hexanchus griseus Fig.2 2b.Snoutlonger,morepointedand narrow;lowerjawwith5rowsof large,comblikeanterolateralteethoneachside;dorsalfinbase separatedfromuppercaudalfin originbyadistancemuchgreater thanitslength.Sizesmaller,upto 1.8 m total length (Fig. 3) - Bigeyesixgill shark ..................................... Hexanchus vituIus 1b.Seven pairs of gill openings (Fig. 4) 3a.Eyesverylarge,headextremely narrowandpointed(Fig.5a); comblikeloweranterolateralteeth longandlow,withashortcusplet anteromedialtothelongmaincusp,andaseriesofabruptly smallercuspletsposterolateraltothecusp(Fig.5b).Bodyplain; withoutspots;sizesmall,toabout 1.4mtotallength(Fig.6)- Sharpnosesevengillshark................Heptranchiasperlo a.underside of headb. tooth Fig. 5 Fig. 4 6 gill openingsHexanchus vitulus Fig. 37 gill openingsHeptranchias perloFig. 6click for next page3b.Eyessmall,headbroadandrounded(Fig. 7a);comblikeloweranterolateralteeth highandshort,withasetofserrations anteromedialtotheshortmaincusp,and aseriesofgraduallysmallercusplets posterolateraltothecusp(Fig.7b).Body usuallywithscatteredsmallblackspots andsometimeswhitespotsalso;size larger,toabout2.9mtotallength(Fig. 8)-Bluntnosesevengillshark............ Notorynchus cepedianus Notorynchus cepedianusFig. 7 Notorynchus cepedianus Fig. 8 Heptranchias Rafinesque, 1810 HEX Hept Genus:HeptranchiasRafinesque,1810,Caratt.gen.sp.anim.piant.Sicilia,Palermo,Pt.1:13.TypeSpecies:SqualuscinereusLacepdebyoriginaldesignation,equalsSqualuscinereusGmelin,1789andajuniorsynonymof Squalus perlo Bonnaterre, 1788. Synonymy :GenusHeptanchusMuller&Henle,1837;GenusHeptranchusGray,1851;GenusHeptrancus Costa, 1857. Diagnostic Features :Headnarrowandpointed,with7pairsofgillslits;eyesverylarge;mouthvery narrowandparabolic;largelowercombliketeethlongandlow,withafewshortmesialcusplets,anabruptlyhigh cusp,andupto7or8distalcuspletsinadults.Caudalpedunclelong,distancefromdorsalfininsertiontoupper caudaloriginovertwicelengthofdorsalfinbase.Colour:spotsabsentfrombody,dorsalfinanduppercaudal lobe wi th black tips, faded or absent in adults but promi nent in young. Remarks:FollowingGarrick&Paul(1971)andBass,d'Aubrey&Kistnasamy(1975c),onlyasinglespeciesis recognized for this genus, H.perlo (Bonnaterre, 1788). Heptranchias perIo (Bonnaterre, 1788) HEX Hept 1SqualusperloBonnaterre,1788,Tabl.encyclop.method.troisreg.Nat.,chthyol.,Paris,10.Holotype: Unknown. Type Locality: Mediterranean Sea. Synonymy :Squalus cinereus Gmelin, 1789;Heptranchias angio Costa, 1857;Notidanus(Heptanchus) cinereus,var.pristiurusvar.aetatis)Bellotti,1877;HeptranchiasdeaniJordan&Starks,1901;Heptranchias dakini Whitley, 1931. a. underside of headb. tooth - 17 -a.click for previous page FAO Names : En - Sharpnose sevengill shark; Fr - Requin perlon; Sp - Canabota bocadulce. upper and lower teeth of left side FieId Marks :Anarrow-headed,big-eyed,smallseven-gilledsharkwith one dorsal fin. Diagnostic Features:See genus Heptranchias. GeographicaI Distribution :Wide-rangingintropicalandtemperateseas;WesternAtlantic:NorthCarolina,USAtoCubaandnorthernGulfofMexico,alsosouthernBrazilandArgentina.EasternAtlantic:FromMoroccotoAngola,alsoMediterraneanSea.ndianOcean:SouthAfrica,southernMozambique,Aldabrasland,southwesternndia.WesternPacific:Japan(southeasternHonshu)andsouthernSeaofJapantoChina,alsondonesia(Bali),Australia(NewSouthWales,Victoria,Tasmania,SouthAustralia)andNewZealand.Eastern Pacific: off northern Chile. Habitat and BioIogy :Marineand benthic,onthecontinentalandinsular shelvesandupperslopes;depthusually between27to720m,butsometimesin shallowerwatercloseinshoreanddownto 1000 m. Aprimaril ydeepwaterspecies,probabl ystrong-swi mming.Ovovivi parous,numberofyoung9to20ina litter.Feedsonbonyfishes,includinghake,andsquid.Veryactiveandaggressivewhencapturedandquickto bite, but too small to be very dangerous to people. Size :Maximumtotallengthabout137cm,sizeatbirthabout26cm,sizeatmaturityabout85cm,for males and 89 to 93 cm for females; said to reach 214 cm, but possibly in error. Interest to Fisheries :Generallycaughtinsomenumbersasabycatchoffisheriesutilizingbottomtrawls and longlines, but of small importance. Literature : Bigelow & Schroeder (1948), Garrick & Paul (1971), Bass, d'Aubrey & Kistnasamy (1975c). Remarks:ThisspecieswasrecentlydiscoveredoffQuilon,southwesternndia(Compagno&Talwar,1982, in press), and off Bali, ndonesia (T. Gloerfelt-Tarp, pers.comm.). - 18 -underside of headclick for next page Hexanchus Rafinesque, 1810 HEX Hex Genus :Hexanchus Rafinesque, 1810, Caratt.gen.sp.anim.piant.Sicilia, Palermo, Pt. 1:14. TypeSpecies: Squalus griseus Lacepde, by original designation, equals Squalus griseus Bonnaterre, 1788. Synonymy :Subgenus Monopterhinus Blainville, 1816 (genus Squalus Linnaeus, 1758);SubgenusNotidanus Cuvier,1817(genusSqualusLinnaeus,1758);GenusHexanchiasSwainson,1838;GenusNotidamusMnster,1842; Genus Hexancus Agassiz, 1846. Diagnostic Features :Headbroadornarrowandpointed,with6pairsofgi llslits;eyessmalltolarge; mouthnarrowandparabolicorwideandarcuate;largelowercombliketeethlowandlong,withmesialserrations,alowtomoderatelyhighcusp,and8to10distalcuspletsinadults;caudalpeduncleshorttolong,distancefrom dorsalfininsertiontouppercaudaloriginvaryingfromslightlylongertoabouttwiceaslongasdorsalbase;body without spots, no black tips on fins. Remarks:FollowingSpringer&Waller(1969),twospeciesarerecognizedforthisgenus,H.griseus (Bonnaterre, 1788) and H. vitulus Springer & Waller, 1969. Hexanchus griseus (Bonnaterre, 1788)HEX Hex SqualusgriseusBonnaterre,1788,Tabl.encyclop.method.troisreg.Nat.,chthyol.,Paris,9.Holotype: Unknown. Type Locality: Mediterranean Sea. Synonymy : SqualusvaccaBloch&Schneider,1801;NotidanusmongeRisso,1826;Hexanchuscorinus Jordan & Gilbert, 1880; Notidanus vulgaris Perez Canto, 1886; Hexanchus griseus australis De Buen, 1960. FAONames:En-Bluntnosesixgillshark;Fr-Requingriset(=Requingris,FishingArea34/47inpart); Sp -