FAO Species Catalogue- Sharks of the world Vol 2 - seafdec.org.my

click for previous pageSharks of the World, Vol. 2 1This is the second volume of an extensively rewritten,revised, and updated version of the original FAO Catalogueof Sharks of the World (Compagno, 1984). It covers all thedescribed species of living sharks of the ordersHeterodontiformes, Lamniformes, and Orectolobiformes,including their synonyms as well as certain well-established butcurrently undescribed species (primarily Australian speciesmentioned by Last and Stevens, 1994). It includes species ofmajor, moderate, minor, and minimal importance to fisheries(Compagno, 1990c) as well as those of doubtful or potential useto fisheries. It also covers those species that have a research,recreational, educational, and aesthetic importance, as well asthose species that occasionally bite and threaten people in thewater and the far more numerous species that are ‘bitten’ andthreatened by people through exploitation and habitatmodification. The Catalogue is intended to form part of acomprehensive review of shark-like fishes of the world in a formaccessible to fisheries workers as well as researchers on sharksystematics, biodiversity, distribution, and general biology. It alsocaters to other researchers that need comparative informationon sharks, people who encounter sharks during the course ofwork or play in the sea or in fresh water, and the general public.This Catalogue builds on a progressive increase in ourknowledge of shark biology over the past two decades, andaddresses an exponential increase in popular interest in sharksand a growing concern over their burgeoning conservationproblems.The term Shark is used here in the broad sense of the FAOInternational Plan of Action for the Conservation andManagement of Sharks (FAO 1999). Sharks include rays orbatoids and chimaeroids as well as ‘nonbatoid’ or ‘typical’sharks, which are the subject of the original shark catalogueand of the present volume. A problem with sharks is thatmost researchers, much less the general public, areunaware of their diversity and tend to focus on the larger,toothy, nonbatoids. There are approximately 1 200 knownliving and valid species of shark-like fishes, cartilaginousfishes, or chondrichthyans, which form the classChondrichthyes. These include at least 50 species of ghostsharks, silver sharks, elephant fish, chimaeras or ratfish(order Chimaeriformes), over 600 species of batoids, flatsharks, or winged sharks (order Rajiformes), and nearly 500species of nonbatoid, ordinary or traditional sharks. Theliving shark-like fishes are included in 10 orders, 60 families,and 186 genera. Diversity of all cartilaginous fishes, livingand extinct, is far greater, with at least 140 valid families, 600valid genera, and at least 3 700 valid species (fromdatabases prepared by the writer).The living cartilaginous fishes are divided into twosister-groups with a long separate, pre-Devonian history, thechimaeroids, Holocephali (with a single living orderChimaeriformes), and the sharks and rays proper orElasmobranchii, with the surviving group subcohortNeoselachii or modern sharks including all of the livingspecies. There is a traditional concept in the taxonomicliterature that divides the living Neoselachii into sharks,Selachii or Pleurotremata, and rays or batoids, Batoidea orHypotremata. Modern cladistic classifications rank thebatoids as an order Rajiformes of the squalomorph sharks(superorder Squalomorphii), and a sister-group of thesawsharks (order Pristiophoriformes) (Fig.1). Hence thebatoids are highly modified, highly diverse, and extremelysuccessful sharks that outnumber all other cartilaginousfishes in species. Chimaeroids are the closest evolutionary1. INTRODUCTIONcousins to elasmobranchs within the Chondrichthyes, andmay find a higher profile as silver sharks or ghost sharks.Considering them as ‘sharks’ brings batoids andchimaeroids out of the perceptual dark. The batoids andchimaeras tend to receive far less attention than nonbatoidsharks in most places. Some of the batoids currently are asimportant for fisheries or more so than nonbatoid sharks orchimaeroids, and some are under severe threat fromoverexploitation and habitat modification (i.e. sawfishes,freshwater stingrays). The batoid sharks will hopefully bethe subject of a forthcoming and much overdue FAOCatalogue of Batoids of the World; likewise for thechimaeroids.Fig. 1 Cladogram showing interrelationships of theorders of living cartilaginous fishesThe original 1984 FAO Shark Catalogue was in one volumein two parts, with pagination across both parts and with asingle bibliography. As the new Catalogue has grown apacewith new information and revisions, it is being published asthree free-standing volumes, each with separate pagination,introduction, terminology, systematic sections, glossary, listof species by FAO Statistical Areas, and a dedicatedbibliography. This will allow readers to independently useeach volume without having to consult the other volumes fortechnical terms and measurements or bibliographicpurposes, as was the case in the old catalogue. We hopethat this added flexibility will be received as an improvement.A larger general introduction to the whole catalogue appearson the first volume and appendices on shark encounters andshark conservation are confined to the third volume.Readers are also encouraged to consult the addendasection included in the last volume of the catalogue. Thepresent and second volume reviews all the species of livingbullhead, mackerel and carpet sharks (ordersHeterodontiformes, Lamniformes and Orectolobiformes),that is, the noncarcharhinoid galeomorph sharks (see Planof the Catalogue below). The first volume covers thenonbatoid squalomorph sharks (orders Hexanchiformes,Squaliformes, Squatiniformes and Pristiophoriformes), andthe third volume reviews the carcharhinoid galeomorphs(order Carcharhiniformes).Apparently sharks are extremely popular at present withconservationists, fisheries managers, the news andentertainment media, and the general public, and are likelyto stay that way for the foreseeable future. Negativeconcepts of sharks were reflected in the 1984 catalogue,sometimes embarrassingly so in hindsight, and partially dueto the negative shark milieu of the times. Hopefully thepresent version departs from this perspective and portrayssharks as a major group of biologically interesting, poorly

2 FAO Species Catalogue for Fishery Purposes No. 1known vertebrates with over 400 million years of unqualifiedsuccess as predators and survivors of mass extinctions.Sharks were then and are now challenged by the ultimateand most terrible of predators, Homo sapiens (‘man, prudentor wise’ as optimistically named by Linnaeus, 1758); butunlike former times the human superpredator is apparentlyaware of the problems and is taking some steps (at last!) tosolve it. One can hope that those efforts are successful.1.1 Plan of the CatalogueThis Catalogue is based on original work on various groupsof sharks as well as my interpretation of data in the literature.Original descriptions of shark species and other taxa wereconsulted if at all possible; when not, various authoritativeworks were consulted for consensus on citations. Some ofthe arrangements of families, genera and species used heredisagree with those of previous workers including those inmy own papers, but in such cases the disagreements arediscussed or reference is made to discussions of suchproblems in the literature. Nonsystematists may notappreciate changes to classification and nomenclaturewrought by systematic studies, and often consider them asannoyances, but shark systematics evolves as does anyother science and changes are inevitable. Hopefully theyare producing increased stability as knowledge improves ina former backwater of systematic ichthyology.Classification and systematic arrangement used here.The present arrangement has evolved from my earlier works(Compagno, 1973, 1977, 1979, 1982, 1984, 1988, 1999),which initially divided the nonbatoid sharks into eight majorgroups or orders and the batoids into four or five orders. Therelationships of the nonbatoid shark orders to one anotherother and to the batoids (order Rajiformes) is approaching atentative consensus following the work of Compagno (1977,1988, 1999 and unpublished), Shirai (1996), and deCarvalho (1996). The following classification of shark-likefishes to order is used in this work and reflects a tentativecladogram based on a summary of previous work andanalysis in progress (* starred orders are covered in thisvolume):Class Chondrichthyes (cartilaginous fishes)Subclass Holocephali (chimaeras and fossil relatives)Order Chimaeriformes (chimaeras or silversharks)Subclass Elasmobranchii (sharks)Cohort Euselachii (modern sharks and fossil relatives)Subcohort Neoselachii (modern sharks)Superorder Squalomorphi (squalomorph sharks)Order Hexanchiformes (cow and frilledsharks)Order Squaliformes (dogfish sharks)Order Squatiniformes (angel sharks)Order Pristiophoriformes (sawsharks)Order Rajiformes (batoids)Superorder Galeomorphi (galeomorph sharks)Order Heterodontiformes (bullhead sharks)*Order Lamniformes (mackerel sharks)*Order Orectolobiformes (carpet sharks)*Order Carcharhiniformes (ground sharks)Orders are the highest taxonomic groups dealt with here,and many of their synonyms are listed even though thepresent International Code of Zoological Nomenclaturedoes not treat groups higher than the family-group level(superfamilies, families, subfamilies, tribes, etc.). Thenomenclature for orders is modified from that of Compagno(1973, 1984, 1999), with synonyms listed from oldest tonewest. The orders are suffixed with -iformes followingcommon ichthyological practice at present. Families aresuffixed with -idae, the universal ending for zoologicalfamilies. Other levels between orders, families, genera andspecies are mostly not covered here. Subgenera arediscussed under their appropriate genera but species arenot grouped under subgenera and given parentheticalsubgeneric names such as Somniosus (Rhinoscymnus)rostratus, even where subgenera are considered valid, soas not to eliminate the utility of listing species alphabeticallywithin genera. Subspecies are listed in the synonymies oftheir species but are not given separate coverage.Valid families, genera and species are provided withcitations for their author or authors, year of publication,reference and pagination (illustrations also included forspecies), while synonyms are similarly cited except for theirreferences (which are listed in the bibliography). Othercombinations of genera and species that have been used inthe literature but are at variance with valid names are citedwith author and date only. The bibliography covers a wideselection of references used in writing the catalogue, but isnot intended to be all-inclusive.The information pertaining to each family, genus andspecies is arranged in the form used in the first edition of thisCatalogue (Compagno, 1984), with some modifications:Family accounts include the valid modern form of thefamily name with author and year; the original citation of thefamily name with its author, year, reference and pagination;the valid type genus with author and date; the number ofrecognized genera in the family; the FAO family vernacularnames in English, French and Spanish; family Synonymswith name, author, year, and pagination; Field Marks andDiagnostic Features of members of the family; an account ofthe natural history of the family under separate sectionscovering Distribution, Habitat and Biology; a section onInterest to Fisheries and Human Impact, a synopsis of thehuman issues affecting shark families; Local Names whenavailable; a Literature section covering references to theentire family; a Remarks section mostly with systematiccomments; and a Key to Genera, when families have morethan one genus.Generic accounts include the valid modern form of thegenus name with author and year; the original citation of thegenus or subgenus, with its author, year, reference andpagination, and, if a subgenus, the original genus name withauthor and year that the subgenus was originally placed in;the type species and means of designating it (for example,by original designation, monotypy, absolute tautonymy, orsubsequent designation); the number of recognized speciesin the genus; the synonyms of genera, with their rank(genus, subgenus, or other genus-group ranking such asW.H. Leigh-Sharpe’s ‘pseudogenera’), author, year,pagination, and genus they were described in if originallyranked as subgenera or equivalents; FAO Names if theyexist; sometimes Field Marks if genera are large anddistinctive; Diagnostic Features of the genus; a Key toSpecies if the genus has more than one species (is notmonotypic); and a Remarks section where necessary.Species accounts include the valid modern names of thespecies, with author and date; the original citation of thespecies (or subspecies), with its author, year, reference

Sharks of the World, Vol. 2 3pagination; the holotype, syntypes, lectotype or neotype ofeach species (paratypes are not listed in the presentaccount), including the total length and sex of the specimen,its institutional deposition, and its catalogue number; thetype locality including the location, coordinates and depth ifavailable, where the holotype, syntypes, lectotype orneotype were caught; Synonyms of the species, includingtheir names, authors and dates; a section listing otherscientific names recently in use; the English, French, andSpanish FAO Names for the species; a lateral viewillustration, and often other useful illustrations (lateral viewdrawings are given of each shark species, usually ventralviews of heads, and often teeth and denticles of the shark inquestion); Field Marks; Diagnostic features (except inmonotypic genera); Distribution, including a map; Habitat;Biology; Size; Interest to Fisheries and Human Impact;Local Names when available; a Remarks section whennecessary; and Literature.Synonyms include only true taxonomic synonyms of thevalid family, genus and species given. For species, anothercategory, Other Combinations, is provided for commonmisidentifications of a given species with another, validspecies (for example, Carcharhinus brachyurus was oftentermed C. remotus, but the latter is a junior synonym of C.acronotus) as well as commonly used combinations thatplace a valid species in different genera (for example,Odontaspis taurus or Eugomphodus taurus for Carchariastaurus).FAO Family and Species Names. English, French andSpanish names for each family and species, primarily foruse within FAO, were selected by the following criteria: (a)each name applies to a single family or species worldwide;(b) the name conforms with FAO spelling nomenclature; (c)the name conforms to prior usage when possible. FAOnames are not intended to replace local species names, butare necessary to overcome the confusion caused by the useof a single name for more than one species or severalnames for one species. The FAO names used here conformwith prior FAO usage and when possible and appropriatenational and international checklists and reviews of speciessuch as Whitley (1940), Fowler (1966-1970), Shiino (1972,1976), Hureau and Monod (1973), Smith (1975), Robins etal. (1980, 1991a, b), and Lindberg, Heard and Rass (1980).The French names were selected jointly with Dr J.C. Quéro,Institut Scientifique et Technique de Pêches Maritimes,Ministère de la Marine Marchande, La Rochelle, France,and for recently discovered species with Dr B. Seret,Museum National d’Histoire Naturelle, Paris. Whenpossible, the names selected correspond to official Frenchspecies nomenclature established by the Direction desPêches Maritimes. The selection of Spanish namespresented considerable difficulties due to the lack ofdenominations for many species. Wherever possible, the“official” Spanish names adopted by F. Lozano in his book“Nomenclature ictiologica”, Madrid, 1963, were used, alongwith names for additional species coined by Dr R. Bonfil,Fisheries Centre, University of British Columbia, Vancouver.The broader use of ‘shark’ here for all living cartilaginousfishes is noted above. The term ‘shark’ is broadly andpopularly used as a catchall term in English for all livingmembers of the Class Chondrichthyes that are not batoidsor chimaeras, although guitarfishes (Rhinobatidae) are alsotermed ‘sand sharks’, chimaeras are termed ‘ghost sharks’or ‘silver sharks’, and even certain aquarium teleosts (someloaches, Cobitidae) are termed ‘sharks’. The French ‘requin’and Spanish ‘tiburón’ are comparable general terms to‘shark’. Several names not incorporating ‘shark’ or itsFrench or Spanish equivalents are mostly used only forsharks and not for other fishes; these include the English‘dogfish’, ‘smoothhound’, ‘tope’, ‘porbeagle’ and‘hammerhead’. However, ‘freshwater dogfish’ is a regionalname for the bowfin, Amia calva, in the USA. ‘Wobbegong’is adapted from an Australian Aboriginal term for sharks ofthe genera Eucrossorhinus, Orectolobus and Sutorectus).French ‘roussette’, ‘emissole’, ‘renard’, ‘milandre’,‘marteau’, and ‘griset’, and Spanish ‘gato’, ‘cazón’, ‘tollo’,‘pintarroja’, ‘tintorera’, and ‘cornuda’, are similar terms forcertain kinds of sharks.Usage of local names for different kinds of sharks variesfrom country to country. ‘Catshark’ is used for members ofthe Scyliorhinidae and sometimes related families (such asProscylliidae) in the United States, but also for variousorectoloboids in Australia. ‘Dogfish’ is variably used formembers of the families Squalidae (‘spiny dogfishes’),Scyliorhinidae (especially Scyliorhinus), and Triakidae(‘smooth dogfishes’, Mustelus spp.). ‘Sand shark’ may referto Odontaspididae (especially Carcharias taurus, the ‘sandtiger shark’ of the eastern USA, called ‘ragged-tooth shark’in South Africa and ‘grey nurse shark’ in Australia), toTriakidae (especially to Mustelus spp.) off the western USA,or guitarfishes off South Africa. In the present Catalogue‘catshark’ is restricted to members of the Scyliorhinidae andProscylliidae (‘false catsharks’ are members of thePseudotriakidae), ‘dogfish’ to the Squaliformes, and ‘sandsharks’ in the form of ‘sand tiger shark’ to theOdontaspididae. Orectoloboid ‘catsharks’ are termed‘carpet sharks’, and ‘sand sharks’ and ‘smooth dogfishes’ ofthe triakid genus Mustelus are termed ‘smoothhounds’(except for M. antarcticus, the Australian ‘gummy shark’).Keys, Field Marks and Diagnostic Features. Thesesections include identification data in different forms. Keys toorders, families, genera and species are standarddichotomous biological keys that are followed in steps ofalternate choices to single out the taxa covered. DiagnosticFeatures are comprehensive lists of characters at theordinal, familial, generic, and species level, with thecharacter choice generally limited to external characters,particularly at the species level, because of spaceconsiderations and their primary purpose of identificationrather than indication of relationships. Some exceptions aretaken with higher taxonomic levels, to support a solid, soundhigher classification. The Diagnostic Features sections arehierarchical, with characters at the ordinal level notduplicated at the family, genus and species levels.Monotypic orders with one family (such asPristiophoriformes), monotypic families with one genus(Chlamydoselachidae) or monotypic genera with onespecies (Carcharodon) all have the Diagnostic Featuressection present only in the highest taxon covered. In amonotypic order, Diagnostic Features are omitted in theaccount of its single family; in a monotypic family, they areomitted from its single genus; and in a monotypic genus,they are omitted from its single species.Field Marks generally include a few obvious characters ofuse in field identification, extracted from DiagnosticFeatures at various levels, but included in a separatesection. Field Marks are listed at the ordinal, familial andspecies levels, and occasionally the generic level in cases oflarge genera with many species. The arrangement of FieldMark characters is semihierarchical and pragmatic and mayinclude characters from a higher level such as an order inlower level taxonomic accounts such as those of species.

Sharks of the World, Vol. 2 5Table 2Limits used to convert upper and lower limits ofdepth ranges (m)For original depth data in the intervalLimit used0-30 031-75 5076-150 100151-250 200251-751 500751 and below 1 000If more than one bathymetric range of distribution wasmentioned (e.g. different ranges for adults and juveniles),the widest range given was used. However, when differentdepth ranges existed for different regions or areas, eachwas chosen and plotted independently.If no depth data was mentioned in the original account,textual descriptions have been translated using the criteriain Table 3 below.Table 3Upper and lower limits of depth ranges (m) used fortextual descriptions of habitatFot text indicatingUpperlimitLowerlimitShelf or continental shelf 0 200Shallow waters, inshore waters, coastal 0 50Continental shelf, neritic 0 200Upper shelf 0 100Deep shelf 100 200Slope 200 1 000Upper slope 200 500Deep slope 500 1 000Terms like benthic, pelagic, surface, bottom deeper water,deepish, great depths included under Habitat were notused. If more than one type of habitat was given, the onecorresponding to the widest possible range of distributionwas used.For species with an oceanic habitat, the main source ofinformation was their known geographic distributionirrespective of depth. Thus, the maps of oceanic speciesgive only information on distribution.All data were transferred to hand drawn maps which weredirectly digitized and georeferenced using WVS andArcWorld for the exact plotting of localities and oceanicprovinces.Where necessary, maps show two different kinds ofdistribution for a given species. Dark red is used to show theknown and certain distribution of a species from reliablerecords, whilst light red or orange is used to show thesuspected or uncertain distribution of a species.Maps presented in the Catalogue can be largely divided intotwo categories, Global (or world maps) and Regional maps.For better visualization, global maps include the speciesdistribution and the land masses especially generalized andprepared from the WVS data set. The regional maps, inaddition to the above, include the 200 m depth isobath as areference of their depth distribution.Note: Whenever the narrowness of the continental or insularshelves and the scale of the maps have caused parts of thedistribution of a species to be undistinguishable, colouredarrows have been used on the map to point to suchdistribution areas.Habitat. Habitat covers information on physical conditionswhere various sharks are found. The known depth range ofthe species (in metres), position in the water column, type ofsubstrate occupied, and preferences relative to coasts arenoted when available. In most cases data on salinity, oxygencontent, and specific temperature of the water in which theyoccur was not available or was not in an easily usable formand has not been regularly compiled here.Biology. Includes data on population structure anddynamics, reproduction, behaviour, sociobiology, age andgrowth, and feeding. Compilation of these data suggeststhat very few species of sharks are biologically well known,and even in the piked dogfish (Squalus acanthias), perhapsthe best-known of living cartilaginous fishes, there are areasof its biology that are very poorly known (such as itsbehaviour and sociobiology). There is a bias in availablenatural history data towards reproductive biology, feeding,and fisheries-related subjects such as age and growth, andcorrespondingly little on ecology, behaviour andsociobiology.Size. All size data are given as total lengths; this is themeasurement most often used as an independent variableand standard measurement in the shark literature, althoughparticularly in fisheries papers precaudal lengths, forklengths, and other measurements have been used fromchoice or necessity. Unfortunately shark workers have notagreed on a standard method of measuring total length, sototal lengths from different sources in the literature may notbe strictly comparable. I prefer and advocate as a standardmethod a direct measurement, in which the shark is heldbelly down with its dorsal caudal-fin lobe depressed into linewith its body axis and total length measured as a point topoint distance (not over the curve of the body) from the snouttip to the tip of the dorsal caudal-fin lobe (see alsoCompagno, 1988). This method lends itself readily to quickuse of a fishboard with a perpendicular front bar or plate toindex the shark’s snout against, a one metre or two metreruler or folding rule slipped under the shark, or even a steelor cloth tape, and avoids the trouble of computation andpossible errors and loss of data.A comparable computational method adding the precaudallength and dorsal caudal-fin margin is advocated bySadowsky (1968). Bigelow and Schroeder (1948) andSpringer (1964) measured total length from the snout tipalong the body axis to a vertical projection from the tip of thedorsal caudal-fin lobe with the caudal fin in a ‘naturalposition’. Bass (1973) advocated a computational methodwhich adds the precaudal length to a number computed bymultiplying the length of the dorsal caudal-fin margin by aconstant (1.0 or less, 0.97 and 0.80 were the numbers) thatcorrects for the different ‘natural angles’ of the caudal axis tothe body axis in different species. The method advocated

6 FAO Species Catalogue for Fishery Purposes No. 1here and in Compagno (1984, 1988) dispenses with allcomputation and avoids arbitrary constants to correct forsupposed ‘natural positions’ of the caudal axis as well as thedifficulties in obtaining accurate vertical projections from thecaudal tips held in ‘natural positions’. Also, with the presentmethod a comparable measurement can be obtained for allor most sharks, rays and chimaeras, although there areproblems with species that have greatly elongatedfilamentous snouts or tails. In contrast methods using‘natural positions’ arbitrarily generate incompatible totallengths for different groups of sharks, and also do not takeinto account changes in the angle of the caudal axis whensharks swim or as they grow (Compagno, 1988).Total length data presented includes maximum size, size atmaturity (in some cases, a size range at maturity, whenabundant data were available) and maximum size for bothsexes (as sexual dimorphism in size is nearly universalamong sharks, with females usually attaining larger sizesthan males, except for some scyliorhinid catsharks wherethe reverse occurs), and size at birth or hatching.Sometimes size at sexual maturity for either or both sexes isnot known, in which cases reported minimum and maximumsizes of adult individuals are given. In some cases maximumsize exceeds that recorded for either sex, in which case thesex of the outsized individual or individuals representing themaximum size measurements was not indicated. In somepoorly known species only immature individuals are known,in which case the hypothetical maximum adult size is almostcertainly larger than the known immature maximum (nocases are known of adult sharks that are considerablysmaller than large immature individuals of the same sex,unlike some other vertebrates). The writer tends to discountold, unverifiable size records of some well-known species,but mentions them as such.Some fisheries biologists and shark researchers useprecaudal length (PCL) or fork length (FL) as standardlengths instead of total length. The first eliminates problemswith sharks having damaged caudal fins but is difficult todetermine on some sharks with weakly defined uppercaudal-fin origins. The second is only applicable to specieswith notched caudal fins and defined upper and lowerpostcaudal-fin margins.In some species length-weight equations are presented,usually of the form W=a+TL b , where W is weight, a and bare constants, and TL is total length.Interest to Fisheries and Human Impact. This section isexpanded in scope from the 1984 catalogue, and in additionto fisheries information includes many other aspects ofhuman interaction with sharks. In this section data onlocalities of fisheries, gear used, and uses of the particularspecies are noted when available. National fisheries data forsharks is often sketchy and combined for a number ofspecies. Thus, catch statistics were available for relativelyfew species of sharks but are noted when available, withparticular emphasis on data from those species reported toFAO. Additional data for sharks are increasingly availablefrom national and regional fisheries bodies, but were used ina very limited way here due to time and literatureconstraints.Initially data from the hard-copy FAO species yearbookswere used for compiling shark fisheries data onspreadsheets, as in Compagno (1990c), but this has beengreatly facilitated by the advent of FAO FishStat, adata-handling and analytical software package which can bedownloaded free from the FAO Fisheries website(http://www.fao.org/fi). FishStat handles a variety ofannually revised FAO fisheries statistics databases and canexport files into other programmes such as spreadsheetsand databases.Conservation and management issues and importance ofsharks to human recreation including ecotouristic diving andvisits to public aquaria are covered in this section. It alsoincludes aspects of shark behaviour that were formerlyplaced in the biology section, that is, shark encounters withpeople. The 1984 Catalogue used the universal term ‘sharkattack’ for encounters when sharks bite or otherwise injurepeople. I have tried to avoid this term in this Cataloguebecause of its extremely negative, subjective, andmisleading connotations, along with a few other hyperbolicterms such as ‘maneater’. I realize that the general publicand especially the news and entertainment media willcontinue to use these emotive terms for a long time despitethe limited realities. It is challenging to think of ways ofdiscussing the subject without the dreadful, gory ‘sharkattack’ image being brought forth, but it does help to buildalternate and more realistic images of a minuscule objectivephenomenon. This is discussed in more detail under SharkEncounters in the third volume of the Catalogue.Local Names. A change from the 1984 Catalogue is thatlocal or regional family and species names in variouslanguages are generally listed when available under aseparate local names heading. These were compiled fromthe same sources used for FAO names (see above), butwhat is presented here is not comprehensive andrepresents what was readily available to the writer. Manyspecies have no vernacular names whatsoever or arelumped under catchall names, while some sharks such asthe white and basking sharks have dozens of names.Obviously some sharks have more of an impact or are muchmore familiar than others, and these get more names (someof which seem like curses or jokes). Wherever possible localnames are presented for important wide-ranging sharks,including fisheries species such as Galeorhinus galeus(‘school shark’ in Australia, ‘tiburón vitaminico’ or ‘vitaminshark’ in Uruguay and Argentina, ‘soupfin’ or ‘oil shark’ offthe Pacific USA and Canada, and ‘vaalhai’ in South Africa)and Carcharias taurus, the very popular shark for fisheries,public aquaria, ecotourism, and conservation (termed‘ragged-tooth shark’ in South Africa, ‘grey nurse shark’ inAustralia, ‘requin sable’ in West Africa, and ‘sand tiger shark’or ‘sand shark’ off the east coast of the United States). Thebroadening interest in sharks and urgent need to acquirespecies-specific data for their management andconservation should encourage fisheries biologists andother researchers to compile local names for their owncountries or regions, and add to our sketchy knowledge oflocal names worldwide.Remarks. Important information, especially on systematicsand nomenclature are given in the remarks section.Literature. References cited here include specific workswith important information for each species and family aswell as comprehensive accounts, but are not intended as acomprehensive bibliography. Reference sections have beenupdated and given more extensive coverage than the 1984Catalogue.

Sharks of the World, Vol. 2 71.2 Technical Terms and Measurements1.2.1 Picture Guide to External Terminology of Sharksfirst dorsal finsnoutnostrileyespiracledorsal-finspinesecond dorsal finprecaudal pitmouthlabialfurrowsgillopeningspelvic finanal finclasper (males)caudalkeelcaudal finpectoral finhead trunk tailFig. 2 Lateral viewnostrilgill slitsprecaudal tailsnoutanal fincaudal fintrunkmouthventpreanal ridgespectoral finpelvic fin(female, no claspers)Fig. 3Ventral viewanterior nasal flap liftedexcurrent aperturenasoral groovemouthsymphisial grooveincurrent apertureincurrentapertureanterior nasal flaplower labial furrowanteriornasal flapcircumnarial groovecircumnarial foldbarbelupper labial furrowexcurrent apertureposteriornasal flapFig. 4 Head of an orectoloboid shark(ventral view)Fig. 5 Nostril

8 FAO Species Catalogue for Fishery Purposes No. 1upper eyelidnotchsecondarylower eyelidnictitatinglower eyelidsubocular pocketlabial furrowFig. 6 Eyeslabial foldspineapexposteriormarginFig. 7 Mouth corneranteriormarginfreereartiporiginbaseinner margininsertion MEDIALANTERIOR LATERAL Fig. 8 Dorsal finepaxial webterminal lobeposterior tipapopylepelvic findorsal lobeterminal margindorsal marginupper originsubterminal marginsubterminal notchclaspergroovelower originupper postventral marginposterior notchhypaxial weblateralfoldclaspershaftpreventral marginlower postventral margincoverrhipidionventral lobeventral tipFig. 9 Caudal finpseudosiphonclasperspurfin insertioninner marginrhipidionclasper glansbasefree rear tiphypopylefin originanteriormarginposteriormarginclasper tipPOSTERIOR Fig. 10 Pectoral finapexFig. 11 Dorsal view of clasper(lamnid shark)

Sharks of the World, Vol. 2 91.2.2 Picture Guide to Skeletal Terminology of SharksANTERIOR rostruminternasal platenasal capsulenasal fontanelesubethmoidfossanasal aperturepreorbital processanterior fontanellecranial roofparietal fossasupraorbital crestorbital notchsuborbital shelfpostorbital processotic capsulebasal platestapedialfenestracarotidforamenforamen magnuma) DORSAL VIEWhyomandibular facetPOSTERIOR occipital centrumb) VENTRAL VIEWrostrumnasalcapsulesupraorbital crestorbitcranial roofotic capsulesphenopterotic ridgerostralnodepterotic horn ANTERIORnasal apertureorbital notchoptic nerve foramenstapedial fenestrasuborbital shelfhyomandibular facetPOSTERIOR c) LATERAL VIEWFig. 12Chondrocraniumbasalsbasalsmetapterygiumpropterygiumradialspropterygiummesopterygiummetapterygiumproximal radialsradialsmetapterygial axismetapterygial axisintermediate radialsdistalradialdistalradialsintermediate radialsa) APLESODIC b) PLESODICFig. 13 Aplesodic and plesodic pectoral fins

Sharks of the World, Vol. 2 111.2.3 Measurements Used for SharksTL = TOTAL LENGTH PP2 = PREPELVIC-FIN LENGTHFL = FORK LENGTH SVL = SNOUT-VENT LENGTHPCL = PRECAUDAL-FIN LENGTH PAL = PREANAL-FIN LENGTHPD2 = PRE-SECOND DORSAL-FIN LENGTH IDS = INTERDORSAL SPACEPD1 = PRE-FIRST DORSAL-FIN LENGTH DCS = DORSAL CAUDAL-FIN SPACEHDL = HEAD LENGTH PPS = PECTORAL-FIN PELVIC-FIN SPACEPG1 = PREBRANCHIAL LENGTH PAS = PELVIC-FIN ANAL-FIN SPACEPSP = PRESPIRACULAR LENGTH ACS = ANAL-FIN CAUDAL-FIN SPACEPOB = PREORBITAL LENGTH PCA = PELVIC-FIN CAUDAL-FIN SPACEPP1 = PREPECTORAL-FIN LENGTH VCL = VENT CAUDAL-FIN LENGTHTLFLPCLPD2PD1HDLPG1PSPIDSPOBDCSPP1PPSPASPCAACSPP2SVLPALVCLFig. 17 Main longitudinal measuresPRN = PRENARIAL LENGTHPOR = PREORAL LENGTHEYL = EYE LENGTHEYH = EYE HEIGHTING = INTERGILL LENGTHGS1 = FIRST GILL SLIT HEIGHTGS2 = SECOND GILL SLIT HEIGHTGS3 = THIRD GILL SLIT HEIGHTGS4 = FOURTH GILL SLIT HEIGHTGS5 = FIFTH GILL SLIT HEIGHTGS6 = SIXTH GILL SLIT HEIGHTGS7 = SEVENTH GILL SLIT HEIGHTP1A = PECTORAL-FIN ANTERIOR MARGINP1R = PECTORAL-FIN RADIAL LENGTHP1B = PECTORAL-FIN BASEP1I = PECTORAL-FIN INNER MARGINP1P = PECTORAL-FIN POSTERIOR MARGINP1H = PECTORAL-FIN HEIGHTP1L = PECTORAL-FIN LENGTHSOD = SUBOCULAR POCKET DEPTHPRNPORSODEYLEYHGS1GS5P1RFig. 18INGP1AP1LP1IP1PP1H

12 FAO Species Catalogue for Fishery Purposes No. 1CDM = DORSAL CAUDAL-FIN MARGINCPV = PREVENTRAL CAUDAL-FIN MARGINCPU = UPPER POSTVENTRAL CAUDAL-FIN MARGINCPL = LOWER POSTVENTRAL CAUDAL-FIN MARGINCFW = CAUDAL-FIN FORK WIDTHCFL = CAUDAL-FIN FORK LENGTHCST = SUBTERMINAL CAUDAL-FIN MARGINCSW = SUBTERMINAL CAUDAL-FIN WIDTHCTR = TERMINAL CAUDAL-FIN MARGINCTL = TERMINAL CAUDAL-FIN LOBECFWCDMCPUCSTCSWCTRCTLD1L = FIRST DORSAL-FIN LENGTHD1A = FIRST DORSAL-FIN ANTERIOR MARGIND1B = FIRST DORSAL-FIN BASED1H = FIRST DORSAL-FIN HEIGHTD1I = FIRST DORSAL-FIN INNER MARGIND1P = FIRST DORSAL-FIN POSTERIOR MARGINCFLCPVCPLD2L = SECOND DORSAL-FIN LENGTHD2A = SECOND DORSAL-FIN ANTERIOR MARGIND2B = SECOND DORSAL-FIN BASED2H = SECOND DORSAL-FIN HEIGHTD2I = SECOND DORSAL-FIN INNER MARGIND2P = SECOND DORSAL-FIN POSTERIOR MARGINFig. 19Measurements of caudal finP2L = PELVIC-FIN LENGTHP2A = PELVIC-FIN ANTERIOR MARGINP2B = PELVIC-FIN BASEP2H = PELVIC-FIN HEIGHTP2I = PELVIC-FIN INNER MARGIN [LENGTH]P2P = PELVIC-FIN POSTERIOR MARGIN [LENGTH]D1AD1HD1PD2LD2B D2IANL = ANAL-FIN LENGTHANA = ANAL-FIN ANTERIOR MARGINANB = ANAL-FIN BASEANH = ANAL-FIN HEIGHTANI = ANAL-FIN INNER MARGINANP = ANAL-FIN POSTERIOR MARGIND1BD1LD1IP2AP2LD2HD2AP2B P2I ANBP2HP2PANAANLANHD2PANIANPFig. 20Measurements of dorsal, pelvic and anal finsPDIPDOHDHTRHABHTAHCPHDPIDPODAO DAIHDH = HEAD HEIGHTTRH = TRUNK HEIGHTABH = ABDOMEN HEIGHTTAH = TAIL HEIGHTCPH = CAUDAL-FIN PEDUNCLE HEIGHTDAI = SECOND DORSAL-FIN INSERTION ANAL-FININSERTIONDAO = SECOND DORSAL-FIN ORIGIN ANAL-FIN ORIGINFig. 21 Other common measurementsDPI = FIRST DORSAL-FIN MIDPOINT PECTORAL-FININSERTIONDPO = FIRST DORSAL-FIN MIDPOINT PELVIC-FIN ORIGINPDI = PELVIC-FIN MIDPOINT FIRST DORSAL-FININSERTIONPDO = PELVIC-FIN MIDPOINT SECOND DORSAL-FINORIGIN

LLASharks of the World, Vol. 2 13MOL = MOUTH LENGTHMOW = MOUTH WIDTHULA = UPPER LABIAL-FURROW LENGTHLLA = LOWER LABIAL-FURROW LENGTHNOW = NOSTRIL WIDTHINW = INTERNARIAL SPACEANF = ANTERIOR NASAL-FLAP LENGTHCLO = CLASPER OUTER LENGTHCLI = CLASPER INNER LENGTHCLB = CLASPER BASE WIDTHCLOCLBNOWANFCLIa) NOSTRILb) CLASPERINWMOLMOWULAc) VENTRAL VIEWd) ANGLE OF MOUTHGIRGIR = GIRTHe) DORSO-LATERAL VIEWSPLINOHDWTRWABWTAWCPWESLf) DORSAL VIEWFig. 22INO = INTERORBITAL SPACESPL = SPIRACLE LENGTHESL = EYE SPIRACLE SPACEHDW = HEAD WIDTHTRW = TRUNK WIDTHABW = ABDOMEN WIDTHTAW = TAIL WIDTHCPW = CAUDAL-FIN PEDUNCLE WIDTH

14 FAO Species Catalogue for Fishery Purposes No. 1Fig. 23 Higher classification of sharks (Orders)

16 FAO Species Catalogue for Fishery Purposes No. 1functionally replaced by expanded neural and haemalarches of the vertebral column.Basal communicating canals: See subnasal fenestrae.Basal groove: In oral teeth, a deep groove proximal to thebasal ledge on the labial surface of the crown neck andapical root margin.Basal ledge: In oral teeth, a shelf-like projection on thelabial surface of the crown foot.Basal plate: The floor of the cranial cavity of theneurocranium, a ventral, medial plate extending from theethmoid region between the orbits and otic capsules andbelow the cranial cavity to the occipital condyles, occipitalcentrum and foramen magnum.Basals or basalia: In a vertebral centrum, the diagonalspaces below the attachment surfaces of the basidorsalcartilages, above the basiventral cartilages, and betweenthe two halves of the double cone. Basalia may be filled withuncalcified cartilage, may have diagonal calcificationspenetrating the uncalcified cartilage, or may have calcifiedannuli or solid calcified cartilage that are continuous withcalcification of the intermedialia. See diagonal calcificationsand intermedialia.Base: In precaudal fins, the proximal part of the fin betweenthe origin and insertion, extending distally, and supported bythe cartilaginous fin skeleton. In the caudal fin, thatthickened longitudinal part of the fin enclosing the vertebralcolumn and between the epaxial and hypaxial lobes or websof the fin. In oral teeth, the proximal root and crown foot, inapposition to the distal cusp. In denticles, the proximalanchoring structures, often with four or more lobes, holdingthe denticles in the skin.Basidorsal cartilages: A pair of wedge-shaped arched, thincartilages articulating with the dorsolateral surfaces of avertebral centrum and forming a continuous neural arch withthe interdorsal cartilages to protect the spinal cord.Basipterygium: The large elongate longitudinal cartilage atthe fin base of the pelvic fin, attached to the posterolateralends of the pelvic girdle or puboischiadic bar. Thebasipterygium has pelvic radials attached along its distaledge and has the clasper skeleton attached posteriorly inmales.Basiventral cartilages: A pair of rounded or wedge-shapedcartilages on the ventrolateral surfaces of a vertebralcentrum that form the bases for attachment of ribs inmonospondylous precaudal vertebrae. In diplospondylousprecaudal and caudal vertebrae the basiventrals formhaemal arches along with the interventral cartilages forprotecting the caudal artery and vein.Batoid: A ray or flat or winged shark, a neoselachian of thesuperorder Squalomorphii, order Rajiformes: a sawfish,sharkray, wedgefish, guitarfish, thornray, panray, electricray, skate, stingray, butterfly ray, eagle ray, cownose ray,devil ray or manta. Rays are closely allied to the sawsharks(Pristiophoriformes) and angel sharks (Squatiniformes), butdiffer from them in having the pectoral fins fused to the sidesof the head over the gill openings, which are ventral ratherthan laterally or ventrolaterally placed.Beta cartilage: In the clasper skeleton, a single,dorsolateral flattened, wedge-shaped or cylindrical cartilageconnecting the pelvic basipterygium and axial cartilage andreinforcing the intermediate segments, possibly derivedfrom a pelvic radial.Blade: In oral teeth, an arcuate, convex-edged section ofthe cutting edge of the crown foot, without cusplets.Body ridges: Elongated longitudinal dermal ridges on thesides of the trunk and precaudal tail in certain carpet sharks(Orectolobiformes), in the whale, zebra and some bamboosharks.Body: Can refer to an entire shark, sometimes restricted tothe trunk and precaudal tail.Branchial arches: The paired visceral arches behind thehyoid arch and just in front of the scapulocoracoid thatsupport the gills. In elasmobranchs the five to sevenbranchial arches primitively consist of a pair of dorsomedialand wedge-shaped cartilages, the pharyngobranchials,closely situated against the roof of the pharynx, a pair ofdorsolateral and more cylindrical epibranchials that areconnected dorsomedially to the pharyngobranchials, a pairof ventrolateral cylindrical ceratobranchials that areconnected ventrolaterally to the epibranchials, a pair ofventromedial hypobranchials that are connectedventrolaterally to the ceratobranchials, and unpairedventromedial basibranchials that are connectedventrolaterally to the hypobranchials. The hypobranchialsand basibranchials along with the expanded ventral ends ofthe ceratobranchials form the basibranchial skeleton ofthe floor of the branchial pharynx. The branchial skeleton isvariably modified in elasmobranchs, with basibranchials andsometimes hypobranchials often lost, the last twopharyngobranchials and the last epibranchial often fusedtogether, and the last basibranchial often expanded into along, broad copula with which the anterior hypobranchialsand posterior ceratobranchials articulate.Calcified cartilage: Shark skeletons are formed of hyalinecartilage or gristle, but this is often reinforced with layers ofcalcified cartilage, cartilage impregnated with a mineral,hydroxyapatite, similar to that of bone but organizeddifferently, in a hard, tile-like pavement of tiny tesserae, ormore compactly as in the calcified structures of vertebralcentra.Calcified double cones: In vertebrae, the primarycalcifications of the notochordal sheath, in lateral viewresembling two hollow, horizontal cones with their apicesmerged, or an hourglass.Cannibal viviparity: See uterine cannibalism.Carcharhinoid: A ground shark, a member of the orderCarcharhiniformes, and including the catsharks, falsecatsharks, finbacked catsharks, barbeled houndsharks,houndsharks, weasel sharks, requiem sharks andhammerheads.Carina: On the crowns of oral teeth, a low blunt mesodistalridge replacing the cusp and cutting edge, in sharks that eathard-shelled invertebrate prey.Carotid foramen: A single foramen or one of a pair offoramina that penetrate the basal plate usually near itsmidlength and allow passage of the internal carotid arteriesinto the cranial cavity. In some advanced elasmobranchs thecarotid foramina shift through the stapedial foramina andonto the medial wall of the orbit.

Sharks of the World, Vol. 2 17Cartilaginous fishes: Members of the classChondrichthyes.Caudal crest: A prominent saw-like row of enlarged pointeddenticles along the dorsal caudal margin and sometimesalong the ventral caudal margin of the caudal fin. Found incertain sharks including hexanchoids and somecarcharhinoids.Caudal fin: The fin on the end of the tail in shark-like fishes,lost in some batoids.Caudal keels: A dermal keel on each side of the caudalpeduncle that may extend onto the base of the caudal fin,and may, in a few sharks, extend forward as a body keel tothe side of the trunk.Caudal peduncle: That part of the precaudal tail extendingfrom the insertions of the dorsal and anal fins to the front ofthe caudal fin.Central foramen: In oral teeth, a nutrient foramen on themidline of the lingual surface of the root, in the transversegroove.Centrum (plural, Centra): A spool-shaped, partially orusually fully calcified structure that forms as a segmentalconstriction in the notochordal sheath of neoselachians, andwhich as an articulated string forms the principal structuralunits of the vertebral column. Centra are primarily formed bythe calcified double cones in the notochordal sheath, whichmay be their only calcification, but additional secondarycalcification may occur in the centrum between the outersurfaces of the calcified double cones, including calcifiedintermedialia, radii, annuli, and diagonal calcifications.Ceratotrichia: Slender soft or stiff filaments of an elasticprotein, superficially resembling keratin or horn, from theGreek keratos, horn, and trichos, hair. Ceratotrichia run inparallel and radial to the fin base and support the fin webs.The prime ingredient of shark-fin soup.Chimaera: A member of the order Chimaeriformes,subclass Holocephali, see also Chimaeroid, Holocephali.Chimaeroid: A chimaera, ratfish, silver shark, ghost shark,spookfish or elephant fish, a member of the orderChimaeriformes.Chondrichthyan: Referring to the class Chondrichthyes.Chondrichthyes: The class Chondrichthyes, from Greekchondros, cartilage, and ichthos, fish, a major taxonomicgroup of aquatic, gill-breathing, jawed, finned vertebrateswith primarily cartilaginous skeletons, 1 to 7 external gillopenings, oral teeth in transverse rows on their jaws, andmostly small, tooth-like scales or dermal denticles.Chondrichthyes include the living elasmobranchs andholocephalans and their numerous fossil relatives, and alsocan be termed shark-like fishes or simply sharks.Chondrocranium: See neurocranium.Circumnarial fold: A raised semicircular, lateral flap of skinaround the incurrent aperture of a nostril, in heterodontoids,orectoloboids, and a few batoids, defined by a circumnarialgroove.Circumnarial groove: A shallow groove defining the lateralbases of the circumnarial folds.Clasper claws: In parascylliid orectoloboids, a longitudinalrow of large anterolaterally directed claw-like denticles onthe dorsolateral surface of the clasper glans, supported bythe terminal ventral.Clasper dactyl: In parascylliid orectoloboids, a largefinger-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 3cartilage of other orectoloboids and other sharks.Clasper gaff or hook: In the external clasper glans, aposterior hook-like structure, like a clasper spur but formedfrom the dorsal terminal cartilage, found in squaloids of thefamily Squalidae.Clasper glans: The distal and dorsal part of the externalclasper from the hypopyle to its tip, and including variousmovable terminal structures; also, the same area of theclasper skeleton.Clasper groove: The longitudinal groove through theclasper, surrounded by the axial and marginal cartilages,and connecting the apopyle and hypopyle.Clasper hooks: In the clasper glans of some carcharhinoidsharks, small claw-like dermal denticles arranged in a rowalong the ventral surface of the free edge of theexorhipidion.Clasper sacs: Dermal sacs with longitudinally ribbed wallson the ventral and medial surfaces of the claspers ofhexanchoids.Clasper shaft: That part of the clasper skeleton from itsorigin on the pelvic fin basipterygium to the hypopyle; also,that part of the external clasper from its base to thehypopyle.Clasper spine: In the external clasper, a projection of theterminal 3 cartilage on the lateral surface of the clasperglans, which forms a short to long, acutely pointed, spinethat is covered with shiny hard tissue, possibly enameloid,dentine or both. In some squaloids other terminal cartilagesmay have spines.Clasper spur: In the external clasper, a projection of theterminal 3 cartilage on the lateral surface of the clasperglans, which may be pointed but is not covered with shinyhard tissue.Clasper tip: The posterior end of a clasper.Claspers: The paired copulatory organs present on thepelvic fins of male cartilaginous fishes, for internalfertilization of eggs, also termed mixopterygia.Classification: The ordering of organisms into groups onthe basis of their relationships, which may be by similarity orcommon ancestry.Cloaca: The common chamber at the rear of the body cavityof elasmobranchs through which body wastes andreproductive products including sperm, eggs, and youngpass, to be expelled to the outside through a commonopening or vent.Cover rhipidion: On the external clasper glans, anelongated, longitudinal blade or flap on its dorsomedial

Sharks of the World, Vol. 2 19have pairs of tendrils or horn-like structures on their ends, orflat flanges on their sides or spiral flanges around theirlengths, which anchor the cases to the bottom. As the eggtravels from the ovaries into the oviducts and through thenidamental glands, the egg case is secreted around it andthe egg is fertilized. Live-bearing sharks may retain eggcases, and these vary from being rigid and similar to those ofoviparous sharks to soft, bag-like, degenerate andmembranous. Soft egg cases may disintegrate during thebirth cycle.Elasmobranch: Referring to the subclass Elasmobranchii.Elasmobranchii: The subclass Elasmobranchii, (fromGreek elasmos, plate, and branchos, gills, in allusion to theirplate-like gill septa), the shark-like fishes other than theHolocephali or chimaeras, and including the living nonbatoidsharks, batoids, and a host of fossil species. They differ fromholocephalans in having 5 to 7 pairs of gill openings open tothe exterior and not covered by a soft gill cover, oral teethseparate and not formed as tooth plates, a fixed first dorsalfin with or without a fin spine, and a short spined or spinelesssecond dorsal.Embryo: An earlier development stage of the young of alive-bearing shark, ranging from nearly microscopic tomoderate-sized but not like a miniature adult. See foetus.Enameloid: The shiny hard external coating of the crownsof shark oral teeth, superficially similar to enamel in landvertebrates.End-style: In the clasper skeleton, the posterior end of theaxial cartilage, between the dorsal and ventral terminalcartilages.Endemic: A species or higher taxonomic group oforganisms that is only found in a given area. It can includenational endemics found in a river system or along part or allof the coast of a given country, but also regional endemics,found off or in adjacent countries with similar habitat, but notelsewhere.Epaxial lobe or web: In the caudal fin, that part of thecaudal fin between the base and dorsal margin, supportedby ceratotrichia.Epaxial web: The entire fin web above the vertebral columnand caudal base.Epiphysial foramen or notch: On the neurocranium, aforamen or notch in the cranial roof at the dorsomedial edgeof the anterior fontanelle, that houses the pineal body.Ethmoid region: That anteriormost sector of theneurocranium including the nasal capsules, internasal platebetween them, and the rostrum.Ethmonuchal muscles: In the orectoloboid familyParascylliidae, paired head muscles that originate on thedorsal myomeres of the nape, and insert via long tendons onthe nasal capsules. These are possibly for elevating thesnout. Not found in any other sharks, though analogousmuscles exist in batoids.Euselachian: Referring to the Euselachii.Euselachii: The cohort Euselachii (Greek Eu, true, good ororiginal, and selachos, shark or cartilaginous fish), thespined or ‘phalacanthous’ sharks, including the modernsharks or Neoselachii, and fossil shark groups including thehybodonts, the ctenacanths, and the xenacanths, allprimitively with anal fins and having two dorsal fins with finspines.Excurrent apertures: The posterior and ventrally facingopenings of the nostrils, which direct water out of the nasalcavities and which are often partially covered by the anteriornasal flaps. These are usually medial on the nostrils andposteromedial to the incurrent apertures, but may beposterior to the incurrent apertures only.Exorhipidion: In claspers, a longitudinally elongated,external blade or flap with its base attached to thedorsolateral edge of the clasper glans, and with its free edgedirected medially. It is supported by the ventral terminal 2cartilage.Eye notch: A sharp anterior or posterior indentation in theeyelid, where present cleanly dividing the upper and lowereyelids.Filter screens: In the whale shark (Rhincodontidae) anddevil rays (Mobulidae), transverse bars with lateral dermallobes on the internal gill openings that form devices forscreening out plankton.Fin skeletons: In unpaired precaudal fins, the basal platesand radials; in the caudal fin, the vertebral column includingexpanded neural and haemal arches; and in the paired fins,the fin girdles, basals, and radials.Fin web: The usually thin, compressed part of the fin, distalto the base, that is supported by ceratotrichia alone (inaplesodic fins) or by ceratotrichia surrounding expanded finradials or by radials only (plesodic fin).First dorsal constrictor muscles: Paired head musclesthat are confluent and functionally part of the levatorpalatoquadrati muscles in most nonbatoid sharks, except inorectoloboids where they are discrete muscles withseparate origins and insertions similar to but more lateralthan the levators.First dorsal fin: The anteriormost dorsal fin of two, rangingin position from over the pectoral fin bases to far posterior onthe precaudal tail.Foetus: A later development stage of the unborn young of alive-bearing shark, that essentially resembles a small adult.Term foetuses are ready to be born, and generally haveoral teeth and denticles erupting, have a colour pattern(often more striking than adults), and, in ovoviviparoussharks, have their yolk-sacs reabsorbed.Foramen magnum: On the neurocranium, the ‘great hole’or posteromedial aperture through the occiput into thecranial cavity, above the occipital centrum and medial andusually dorsal to the occipital condyles. The spinal cordpasses from the brain through the foramen magnum into theneural canal of the vertebral column.Free rear tips: The pectoral, pelvic, dorsal, and anal fins allhave a movable rear corner or flap, the free rear tip, that isseparated from the trunk or tail by a notch and an innermargin. In some sharks the rear tips of some fins are veryelongated.

20 FAO Species Catalogue for Fishery Purposes No. 1Frontal and parietal fenestrae: On the neurocranium,medial apertures in the cranial roof between the anteriorfontanelle and the parietal fossa, the frontal fenestra beingcloser to the anterior fontanelle and the parietal fenestra tothe parietal fossa. Sometimes the two merge and become afrontoparietal fenestra, while in many batoids and in someorectoloboid sharks there is a merging of the anteriorfontanelle with the frontoparietal fenestra so that it extendsnearly to the parietal fossa. All of these fenestrae are closedby tough membranes.Functional series: A series of oral teeth that are infunctional position on the jaw.Galeomorph: Referring to the Galeomorphii.Galeomorphii: The neoselachian superorderGaleomorphii, including the heterodontoid, lamnoid,orectoloboid, and carcharhinoid sharks.Gill openings or slits: In elasmobranchs, the paired rows offive to seven transverse openings on the sides or undersideof the head for the discharge of water through the gills.Chimaeras have their four gill openings hidden by a soft gillcover and discharge water through a single external gillopening.Gill-raker denticles: In the basking shark (Cetorhinidae),elongated denticles with hair-like cusps arranged in rows onthe internal gill openings, which filter out planktonicorganisms.Gill-raker papillae: Sparse to dense dermal papillae on thegill arches of some sharks that serve as filters to collectsmall food organisms.Girdle: A bar of cartilage buried in the body wall thatsupports the basals of the paired fins: the pectoral girdle(scapulocoracoid) and pelvic girdle (puboischiadic bar).Haemal arch: The arch ventral to the notochord or vertebralcentra on tail vertebrae that is formed by the basiventralsand interventrals and which houses the caudal artery andcaudal vein in a haemal canal.Haemal spines: On the haemal arches of thediplospondylous precaudal and caudal vertebrae, elongatedventral surfaces forming vertical plates, particularlywell-developed on the caudal fin.Head: That part of a cartilaginous fish from its snout tip tothe last or (in chimaeras) only gill slits.Heterocercal: A caudal fin with the vertebral axis slanteddorsally into the fin base, which is also dorsally elevated.Heterodontoid: A bullhead shark, horn shark, or PortJackson shark, a member of the order Heterodontiformes,family Heterodontidae.Heterodonty: In oral teeth, structural differences betweenteeth in various positions on the jaws, between teeth in thesame position during different life stages, or between teethin the same positions in the two sexes.Hexanchoid: A cowshark or frilled shark, members of theorder Hexanchiformes, and including the sixgill sharks,sevengill sharks, and frilled sharks.Holocephali: The subclass Holocephali (from Greek holos,entire, and kephalos, head), the living chimaeras and theirnumerous fossil relatives, a major subdivision of the classChondrichthyes. The name is in reference to the fusion ofthe upper jaws or palatoquadrates to the skull in all livingspecies and in many but not all fossils. The livingholocephalans include three families in the orderChimaeriformes. The living species differ fromelasmobranchs in having four pairs of gill openings coveredby a soft gill cover and with a single pair of external gillopenings, oral teeth fused and reduced to three pairs ofever-growing tooth plates, an erectile first dorsal fin with aspine and a long, low spineless second dorsal.Holotype: Either the only specimen used and mentioned inan original description of a species, with or without adesignation of such, or one of two or more specimens usedand mentioned in an original description of a species anddesignated as such. This becomes the ‘name-bearer’ of thespecies, and is used to validate the species or scientificname by anchoring it to a single specimen.Homodonty: In oral teeth, structural similarity betweenteeth in various positions on the jaws, between teeth in thesame position during different life stages, or between teethin the same positions in the two sexes.Hyoid arch: The visceral arch that supports the tongue and,in elasmobranchs, the rear of the upper jaws. The hyoid archis between the mandibular arch and the first branchial arch,and has the spiracular pocket between it and the mandibulararch. The hyoid arch in elasmobranchs includes a medialbasihyoid in the floor of the mouth and inside the tongue, apair of elongated ceratohyals articulating with the basihyoidand the hyomandibulae, and a pair of hyomandibulaearticulating with the ceratohyals and the hyomandibularfacets of the neurocranium. Chimaeroids have anonsuspensory hyoid arch similar to the gill arches, with apair of epihyals and pharyngohyals equivalent to thehyomandibulae. Batoids have the ceratohyals reduced andseparated from the hyomandibulars or absent, andfunctionally replaced by paired dorsal and ventralpseudohyoids.Hyomandibular facet: On the neurocranium ofelasmobranchs, a joint surface, socket or cotyle that isusually on the ventrolateral surfaces of each otic capsule butmay be extended posteriorly or arched dorsally. The headsof the hyomandibulae articulate with these facets.Chimaeras lack hyomandibular facets and differentiatedhyomandibulae.Hyomandibular nerve foramina: Foramina for the roots ofthe hyomandibular nerves, behind the orbital fissures.These foramina are confluent with the orbital fissure in manysharks.Hypaxial web: The entire fin web below the vertebralcolumn (vertebral axis) and the caudal base.Hypercalcified structures: Parts of the skeleton that havedeveloped extremely dense calcified cartilage, primarilyduring growth and maturation, which sometimes swell toknobs that distort and engulf existing cartilaginousstructures. The rostrum of the salmon shark (Lamnaditropis) is a particularly impressive hypercalcifiedstructure.Holocephalan: Referring to the Holocephali.

Sharks of the World, Vol. 2 21Hypocercal: A caudal fin with the vertebral axis slantedventrally into the fin base, which is also ventrally depressed.Found only in angel sharks (Squatiniformes) among livingsharks.Hypopyle: On the external clasper and clasper skeleton,the posterior opening of the clasper groove onto the clasperglans.Incurrent apertures: The anterior and ventrally facingopenings of the nostrils, which direct water into the nasalcavities. These are usually lateral on the nostrils andanterolateral to the excurrent apertures, but may be anteriorto the excurrent apertures only.Independent dentition: Teeth along a mesodistal series inwhich the roots do not overlap and are separated by aspace. See overlapping dentition.Inner margin: In precaudal fins including the pectoral,pelvic, dorsal and anal fins, the margin from the fin insertionto the rear tip.Insertion: The posterior or rear end of the fin base inprecaudal fins. The caudal fin lacks insertions except withmany batoids and some chimaeroids that have a caudalfilament that extends posterior to the fin. See origin.Interdorsal cartilages: A pair of wedge-shaped arched thincartilages fitting between the basidorsal cartilages of eachvertebra to complete the neural arch.Interdorsal ridge: A ridge of skin on the midback of sharks,in a line between the first and second dorsal fins; particularlyimportant in identifying grey sharks (genus Carcharhinus,family Carcharhinidae).Intermedialia: In a vertebral centrum, dorsal, ventral andlateral spaces between the attachment surfaces of thebasidorsal and basiventral cartilages and between the twohalves of the double cone. These can be filled withuncalcified cartilage, with solid or hollow wedges of calcifiedcartilage, or with plate-like, branched calcified radii withinuncalcified cartilage. See basalia.Intermediate segments: In the clasper skeleton, one ormore short cylindrical cartilages connecting the pelvicbasipterygium to the axial cartilage of the clasper. Alsotermed stem-joints.Intermediate teeth: Small oral teeth between the lateralsand anteriors of the upper jaw, found in most lamnoids.Internasal plate or septum: On the neurocranium, a plateor partition between the two nasal capsules. It ranges from avertical plate to a broad horizontal plate.Interventral cartilages: A pair of rounded or wedge-shapedcartilages fitting between the basiventral cartilages of eachvertebra, that in diplospondylous precaudal and caudalvertebrae form the haemal arches with the basiventralcartilages.Intestinal valve: A dermal flap inside the intestine,protruding into its cavity or lumen, and of various forms indifferent cartilaginous fishes. Often formed like a corkscrewor augur. See spiral, ring and scroll valves.Jaws: See mandibular arch.Labial cartilages: Paired cartilages that are internal andsupport the labial folds at the lateral angles of the mouth.Living neoselachians typically have two pairs of upper labialcartilages, the anterodorsal and posterodorsal labialcartilages, and one pair of ventral labial cartilages, butthese are variably reduced and sometimes absent in manysharks. Chimaeras have more elaborate labial cartilagesthan living elasmobranchs.Labial flange: On tooth crowns of many squaloids andsome orectoloboids, a narrow, vertically elongated labialbasal ledge.Labial folds: Lobes of skin at the lateral angles of themouth, usually with labial cartilages inside them, separatedfrom the sides of the jaws by pockets of skin (labial groovesor furrows).Labial furrows or labial grooves: Grooves around themouth angles on the outer surface of the jaws of manycartilaginous fishes, isolating the labial folds. Primitivelythere is a distinct upper labial furrow above the mouthcorner and a lower labial furrow below it.Labial: In oral teeth, the outer face of the tooth that isdirected outside the mouth and towards the lips. Seelingual.Lamnoid: A mackerel shark, a member of the orderLamniformes, and including the sand tiger sharks, goblinsharks, crocodile sharks, megamouth shark, threshersharks, basking shark, and the makos, porbeagle, salmonshark and white shark.Lateral clasper fold: In mackerel sharks (family Lamnidae),a unique longitudinal flap of skin along the lateral edge of theexternal clasper shaft.Lateral commissures: On the neurocranium, tube-like orring-like enclosed passages for the lateral head veins, whichdrain the orbital sinuses, through the postorbital walls of theorbits and below the sphenopterotic ridges and above thehyomandibular facets in neoselachians. The lateralcommissures are reduced or absent in many livingneoselachians.Lateral or laterad: Outwards, in the transverse directiontowards the periphery of the body. See medial.Lateral orolabial grooves: Shallow longitudinal grooves onthe lower jaw that connect the edge of the lip on each sidewith the medial ends of the lower labial furrows. Found inmore advanced orectoloboids.Lateral teeth: Large broad-rooted, compressed, highcrowned oral teeth on the sides of the jaws between theanteriors and posteriors.Lateral trunk denticle: A dermal denticle from thedorsolateral surface of the back below the first dorsal finbase.Lectotype: One of two or more specimens that weresyntypes in an original description, designated as alectotype by a subsequent writer. It then becomesequivalent to a holotype, and anchors the name of thespecies to a specimen unless invalidated by a ruling of theInternational Commission on Zoological Nomenclature or aprevious designation of a lectotype.

22 FAO Species Catalogue for Fishery Purposes No. 1Levator palatoquadrati muscles: Paired head musclesthat primitively originate on the underside of the postorbitalprocesses and sphenopterotic ridges, extend vertically, andinsert on the posteromedial surfaces of the quadrateprocesses of the palatoquadrates. In advancedcarcharhinoids the origins of the levator palatoquadratimuscles are expanded far forwards and diagonally into theorbits. Primitively these muscles lift or retract the jawsupwards, but in advanced carcharhinoids may help rotatethe jaws forwards and downwards in opposition to thelevator hyomandibularis muscles, which retract the jaws.Lingual: In oral teeth, the inner face of the tooth that isdirected inside the mouth and towards the tongue. Seelabial.Live-bearing: A mode of reproduction in which femalesharks give birth to young sharks, which are miniatures ofthe adults. See viviparity.Longitudinal ridges: In lateral trunk denticles, parallelridges that extend anteroposteriorly on the distal surface ofthe crown. These may be in the form of a single medialridge (sometimes paired), and paired lateral ridges, andmay terminate in medial and lateral cusps.Lower eyelid: The ventral half of the eyelid, separated by adeep pocket (conjunctival fornix) from the eyeball. In somederived batoids the pocket also fuses with the eyeball.Lower origin: In the caudal fin, the anteroventral beginningof the hypaxial or lower web of the caudal fin, at the posteriorend of the anal-caudal or pelvic-caudal space (seemeasurement illustrations).Lower postventral margin: In the caudal fin, the lower partof the postventral margin of the hypaxial web, from theventral tip to the posterior notch.Mandibular arch: The paired primary jaw cartilages ofsharks, including the dorsal palatoquadrates and the ventralMeckel’s cartilages.Mandibulocutaneous muscles: Paired head muscles insqualoid and hexanchoid sharks, that originate on the insideof the skin of the head behind the eyes and near thespiracles, and insert on the dorsoposterolateral face of thequadrate processes of the palatoquadrates.Meckel’s cartilages: The paired lower jaw cartilages,articulating mesially with each other at the midline orsymphysis of the lower jaw, and articulating laterally with thedistal ends of the palatoquadrates. The Meckel’s cartilagesare fused together at the symphysis in some shark-likefishes or are articulated to a symphysial cartilage in others.Medial teeth: Small oral teeth, generally symmetrical andwith narrow roots, in one row at the symphysis and often inadditional paired rows on either side of the symphysial one.Medial: Inwards, in the transverse direction towards themiddle of the body. See lateral.Mesial: In oral teeth, mesial structures are towards themidlines of the jaws, the symphyses. See distal.Mesopterygium: In the pectoral fin skeleton of livingneoselachians, the middle basal cartilage, between thepropterygium and metapterygium. The mesopterygium issometimes fused to the propterygium or metapterygium, orto both.Mesorhipidion: A knife-like or blade-like structure on thelateral clasper glans of some carcharhinoid sharks, formedfrom the terminal 3 cartilage, and over and partially lateral tothe ventral terminal and mesial to the pseudopera.Metapterygial axis: In the pectoral fin skeleton of livingneoselachians, the posterior extension of themesopterygium as a flattened, elongated segmented seriesof cartilages that supports the distal bases and free rear tipsof the pectoral fins; the axis has radials along its distal edgecontinuous with the radials on the metapterygial basal.Metapterygial basal: In the pectoral fin skeleton of livingneoselachians, the anteriormost, expanded cartilage of themetapterygium.Metapterygial proximal segment: In the hexanchoidpectoral fin skeleton, a short jointed segment on theproximal end of the metapterygial basal, not found in othersharks.Metapterygium: In the pectoral fin skeleton of livingneoselachians, the rearmost basal cartilage, adjacent to theposterior edge of the mesopterygium and with severalradials attached to its distal edge. It includes themetapterygial basal and the metapterygial axis.Molariform: In oral teeth, referring to a tooth with a broadflat crown with low cusps or none, for crushing hard-shelledinvertebrate prey.Monospondylous precaudal vertebrae: Vertebrae withone centrum and one pair of basidorsals, basiventrals, andribs per body segment (myotome), and generally extendingfrom the occiput to the end of the body cavity and to over thepelvic girdle. However there is much variation in the positionof the monospondylous-diplospondylous transition, whichcan range well in front or behind the pelvic girdle.Monospondylous-diplospondylous transition: Theposition on the vertebral column where monospondylouscentra end and diplospondylous centra begin. In lateral viewthe transition often appears as an abrupt decrease in lengthof the diplospondylous centrum compared to the lastmonospondylous centrum, but this can be obscure invarious sharks with very numerous, very short centra. Oftena centrum of intermediate length appears between a longmonospondylous centrum and a short diplospondylouscentrum. In a few sharks there is a stutter zone ofalternating long and short centra that marks the transition.Also, the basidorsals and basiventrals have foramina for thespinal nerves on every other vertebra, rather than on eachvertebra as in monospondylous vertebrae. The transitionfrom long to short centra is generally coordinated with thetransition of vertebrae with free ribs and no haemal arches tothose without ribs and with haemal arches. However, insome sharks the two transitions can be anterior or posteriorto each other.Multiple oviparity: A mode of egg-laying or oviparity inwhich female sharks retain several pairs of cased eggs inthe oviducts, in which embryos grow to advanceddevelopmental stages. When deposited on the bottom (incaptivity) the eggs may take less than a month to hatch.Found only in the scyliorhinid genus Halaelurus, with someuncertainty as to whether the eggs are normally retained in

Sharks of the World, Vol. 2 23the oviducts until hatching. Eggs laid by these sharks maybe abnormal, unusual, or an alternate to ovoviviparity. Thewhale shark (Rhincodon typus) may have multiple retentionof egg cases; near-term foetuses have been found in theiruteri and egg-cases with developing foetuses have beencollected on the bottom.Nasal aperture: On the neurocranium, an aperture in theanteroventral surface or floor of each nasal capsule, throughwhich the nostril directs water into and out of the nasalorgan.Nasal capsules: On the neurocranium, a pair of spherical,oval or trumpet-shaped, thin-walled structures behind therostrum (when present) and in front of the orbits, cranial roofand basal plate. They serve as containers for the nasalorgans or organs of smell, and have passages into thecranial cavity to connect the nasal organs with the brain.Nasal curtain: Anterior nasal flaps that are expandedmedially and posteriorly and have fused with each other.Nasal curtains are found in some carcharhinoid sharks andin many batoids.Nasal flap: One of a set of dermal flaps associated with thenostrils, and serving to direct water into and out of them,including the anterior, posterior, and mesonarial flaps.Nasal fontanelle: On the neurocranium, an aperture in theposteroventral surface or floor of each nasal capsule,behind the nasal apertures and closed by a dermalmembrane.Nasoral grooves: Many bottom-dwelling, relatively inactivesharks have nasoral grooves, shallow or deep grooves onthe ventral surface of the snout between the excurrentapertures and the mouth. The nasoral grooves are coveredby expanded anterior nasal flaps that reach the mouth, andform water channels that allow the respiratory current to pullwater by partial pressure into and out of the nostrils and intothe mouth. This allows the shark to actively irrigate its nasalcavities while sitting still or when slowly moving. Nasoralgrooves occur in heterodontoids, orectoloboids,chimaeroids, some carcharhinoids, and most batoids. Alsotermed oronasal grooves.Neck: A narrow band of finely porous dull tissue (possiblyorthodentine) encircling the proximal end of the crown of atooth, and apparently covered with dental membrane.Neoselachian: Referring to the Neoselachii.Neoselachii: From Greek neos, new, and selachos, shark.The modern sharks, the subcohort Neoselachii, consistingof the living elasmobranchs and their immediate fossilrelatives. See Euselachii.Neotype: A specimen, not part of the original type series fora species, which is designated by a subsequent author,particularly if the holotype or other types have beendestroyed, were never designated in the original description,or are presently useless.Neural arch: In shark vertebrae, a dorsal arch formed bybasidorsal and interdorsal cartilages above the centrum andforming a neural canal containing the spinal cord.Neural spines: On the neural arches of shark vertebrae,elevated dorsal plate-like surfaces, particularlywell-developed in many squalomorph sharks.Neurocranium: In sharks, a box-shaped complexcartilaginous structure at the anterior end of the vertebralcolumn, containing the brain, housing and supporting thenasal organs, eyes, ears, and other sense organs, andsupporting the visceral arches or splanchnocranium. Alsotermed chondrocranium, chondroneurocranium, orendocranium.Nictitating lower eyelid: In the ground sharks (orderCarcharhiniformes), a movable lower eyelid that has specialposterior eyelid muscles that lift it and, in some species,completely close the eye opening (or palpebral aperture).Often incorrectly termed nictitating membrane, a different,nonhomologous structure in terrestrial vertebrates.Nictitating upper eyelid: In parascylliid orectoloboids, theupper eyelid has anterior eyelid muscles that pull it downand close the eye opening, analogous to the nictitating lowereyelids of carcharhinoids.Nomenclature: In biology, the application of distinctivenames to groups of organisms.Nostrils: The external openings of the cavities of the nasalorgans, or organs of smell.Notochord: In embryonic sharks (and other chordates) thenotochord is a fluid-filled tube below the spinal cord that hasa connective-tissue notochordal sheath surrounding it. Thenotochord forms the primitive developmental base of thechondrichthyan vertebral column. Chimaeroids retain thenotochord and its sheath without constriction (althoughsome have ring-like centra in the sheath), but inneoselachians it is constricted by the development ofdouble-cone calcifications of the centra within the sheathinto biconical chambers between each centrum. Theaddition of centra to the notochordal sheath strengthens thevertebral column. Some deepwater squaloid, hexanchoid,and lamnoid sharks have the sheath constriction andcalcified double cones variably reduced, sometimes toconnective tissue septa only. Some of these taxa with a‘notochordal’ vertebral column have been consideredprimitive but are apparently derived from ancestors withwell-calcified, constricted vertebral centra.Occipital centrum: On the occiput of the neurocranium, theposterior half of a calcified double cone of the vertebralcolumn, imbedded in the basal plate and articulating with theanteriormost centrum of the vertebral column. Also termedoccipital hemicentrum.Occiput: The posteriormost sector of the neurocranium,behind and partially between the otic capsules, with itsdorsal surface from the parietal fossa rearwards to theforamen magnum, and its posterior surface including theoccipital condyles, the occipital centrum, the paired vagusnerve foramina, the paired glossopharyngial nerveforamina, and the rear surface of the hyomandibular facets.Ocelli or eyespots: Large eye-like pigment spots locatedon the dorsal surface of the pectoral fins or bodies of somesharks including rays, angel sharks, and some bamboosharks, possibly serving to frighten potential enemies.

24 FAO Species Catalogue for Fishery Purposes No. 1Oophagy: From Greek oön, egg, and phagos, to eat.Egg-eating, a mode of live-bearing reproduction employinguterine cannibalism; early foetuses deplete their yolk-sacksearly and subsist by eating nutritive eggs produced by themother. Known in several lamnoid sharks, the carcharhinoidfamily Pseudotriakidae, and in the orectoloboid familyGinglymostomatidae (Nebrius ferrugineus).Optic nerve foramen: A large foramen usually in the middleof the orbital wall, passing the optic nerve from the brain tothe eye.Optic pedicel: On the neurocranium, a slender cartilagethat projects from the medial orbital wall and articulates withthe eyeball; it serves as a pivot point for the eyeball and aspacer between the eyeball and the orbital wall.Orbital fissures: The main foramina or fenestrae that passthe trigeminal and facial nerves from the brain to the orbits,located on the posteroventral ends of the medial walls of theorbits.Orbital notches: On the neurocranium, the paired anteriornotches in the suborbital shelves that articulate with theorbital processes of the palatoquadrates. In manysqualomorph sharks these are enlarged, deepened,socket-like, and posteriorly situated in the orbits, withtelescoping of the suborbital shelves, and are lost in batoids.Orbits: Large, paired cavities on the sides of theneurocranium, behind the nasal capsules, mostly in front ofthe otic capsules, and separated medially by the cranialcavity. They are bounded anteriorly by the preorbital wallsand processes, dorsally by the supraorbital crests, ventrallyby the suborbital shelves (reduced or lost in varioussqualomorphs), and posteriorly by the postorbital processesand walls. The orbits contain the eyeballs and their muscles,venous sinuses, several arteries that connect to the cranialcavity, and most of the cranial nerves.Orectoloboid: A carpet shark, a member of the orderOrectolobiformes, including barbelthroat carpet sharks,blind sharks, wobbegong sharks, bamboo sharks, epaulettesharks, nurse sharks, zebra sharks, and whale sharks.Origin: The anterior or front end of the fin base in all fins.The caudal fin has upper and lower origins but no insertion.See insertion.Orthodentine: A primary hard tissue comprising the crownof oral teeth in sharks, with numerous fine mostly parallelnonvascular tubules.Orthodont: An oral tooth with its crown filled withorthodentine, and with a prominent central pulp cavity.Osteodentine: A primary hard tissue comprising the rootsand sometimes the inside of the crown in the oral tooth, withbone-like large reticulating, thick-walled tubules.Osteodont: An oral tooth with its crown filled withosteodentine, continuous with the root, and without a pulpcavity.Otic capsules: On the neurocranium, a pair of complexthick-walled capsules containing the inner ears, and locatedbetween the orbits and the occiput, and partially separatedmedially by the cranial cavity.Overlapping dentition: Teeth along a mesodistal series inwhich the roots overlap and are not separated by a space.Two types of overlap patterns occur, alternate overlap, inwhich teeth in a series alternate from more labial to morelingual, and imbricate overlap, in which the distal end ofeach tooth lingually or labially overlaps the mesial end of thesucceeding tooth, repeating to the distal ends of the dentalband. Alternate-imbricate dentitions combine bothalternate and imbricate overlap. See independentdentition.Oviparity: A mode of reproduction in which female sharksdeposit eggs enclosed in oblong or conical egg-cases on thebottom, which hatch in less than a month to more than ayear, producing young sharks which are miniatures of theadults.Ovoviviparity: Generally equivalent to yolk-sac viviparity,live-bearing in which the young are nourished primarily bythe yolk in the yolk-sac, which is gradually depleted and theyolk-sac reabsorbed until the young are ready to be born.Sometimes used to cover all forms of aplacental viviparity,including cannibal viviparity.Paired fins: The pectoral and pelvic fins.Palatoquadrates: The paired upper jaw cartilages,articulating mesially with each other at the midline orsymphysis of the upper jaw, and articulating laterally with thedistal ends of the Meckel’s cartilages. The palatoquadratesare fused to the neurocranium in all living holocephalans.The palatoquadrates of neoselachians are divided intocylindrical anteromedial sectors or palatine processes,which articulate or are otherwise attached to each other atthe symphysis; variably modified conical to flattenedarticular structures or orbital processes on the middle ofthe palatoquadrates for attachment to the neurocranium atthe orbital notches; and often elevated posterodistalquadrate processes that articulate with the distal ends ofthe Meckel’s cartilages and are loosely or firmly attached tothe distal ends of the hyomandibulae. In a few livingneoselachians, and many fossil elasmobranchs, thequadrate processes have postorbital articulations withthe rear surfaces of the postorbital processes of theneurocranium.Palpebral aperture: The eye opening, defined by the upperand lower eyelids.Papillae: Elongated finger-like processes of skin, locatedaround the spiracles of torpedo rays, and in the mouths andon the gill arches of other sharks.Papillose gill rakers: See gill raker papillae.Paralectotype: One of two or more specimens that weresyntypes in an original description, but which became aparalectotype or paralectotypes when a subsequent authordesignated one of the syntypes as a lectotype.Paralectotypes are equivalent to paratypes.Paratype: Each specimen of a type series other than theholotype. Specimens other than the holotype automaticallybecome paratypes unless the author designates them asreferred specimens that are not part of the type series.Parietal fossa: On the neurocranium, a shallow or deepdepression between the otic capsules and at the rear of the

Sharks of the World, Vol. 2 25cranial roof, that houses foramina for paired ducts leading tothe inner ears and for the spaces around them.Pectoral fins: A symmetrical pair of fins on each side of thetrunk just behind the head and in front of the abdomen.These are present in all cartilaginous fishes and correspondto the forelimbs of a land vertebrate (a tetrapod orfour-footed vertebrate).Pectoral or shoulder girdle: See scapulocoracoid.Pedicel: In lateral trunk denticles, a narrow stalk separatingthe crown from the base.Pelvic fin: A symmetrical pair of fins on the sides of the bodybetween the abdomen and precaudal tail which correspondto the hindlimbs of land vertebrate (a tetrapod or four-footedvertebrate). Also, ventral fins.Pelvic girdle: See puboischiadic bar.Photophores: Conspicuously pigmented small spots on thebodies of most lantern sharks (family Etmopteridae) andsome kitefin sharks (family Dalatiidae). These are tiny roundorgans that are covered with a conspicuous dark pigment(melanin) and produce light by a low-temperature chemicalreaction.Placenta: See yolk-sac placenta.Placental viviparity: Live-bearing in which the youngdevelop a yolk-sac placenta, which is apparently confined tothe carcharhinoid sharks.Placoid scale: See dermal denticle.Plesodic fin: A pectoral, pelvic, dorsal, or anal fin in whichthe radial cartilages of the fin skeleton extend far into thedistal fin web, often near its edges, and between thesupporting ceratotrichia of the fin web. Some fossil sharksalso have plesodic caudal fins, in which the expandedhaemal arches of the caudal vertebrae extend far into the finweb. In more advanced batoids the radials of the plesodicpaired fins become highly branched and segmented, verynarrow and slender, and essentially replace the ceratotrichiaas supports for the fin webs.Pores, pigmented: In a few sharks and skates, the poresfor the lateral line and ampullae of Lorenzini areconspicuously black-pigmented, and look like little blackspecks.Posterior: Rearwards, in the longitudinal direction of thecaudal-fin tip or tail filament. Also caudal.Posterior margin: In precaudal fins, the margin from the finapex to either the free rear tip (in sharks with distinct innermargins) or the fin insertion (for those without innermargins).Posterior nasal flaps: Low flaps or ridges arising on theposterior edges of the excurrent apertures of the nostrils.Posterior notch: In the caudal fin, the notch in thepostventral margin dividing it into upper and lower parts.Posterior teeth: Small or sometimes enlarged irregular oralteeth near and at the distal ends of the dental bands, withlow crowns and sometimes missing cusps.Posterior tip: The posteriormost corner or end of theterminal lobe of the caudal fin.Postocular eyelid muscles: A complex of paired headmuscles unique to carcharhinoid sharks that originatearound the spiracles and insert on the posterior ends of theupper eyelids and nictitating lower eyelids. Primitively theydepress the upper eyelid and elevate the nictitating lowereyelid to close the eye, but in more derived carcharhinoidsthe eye is closed only by elevation of the nictitating lowereyelid.Postorbital processes: On the neurocranium,posterolateral projections of the supraorbital crests, belowwhich the postorbital walls originate.Postorbital walls: On the neurocranium, the posteriorboundaries of the orbits, variously reduced vertical plates ofcartilage that close the orbits between the postorbitalprocesses and the suborbital shelves, more or less reducedin living neoselachians.Postventral margin: In the caudal fin, the margin from theventral tip to the subterminal notch of the caudal fin. Seelower and upper postventral margins.Preanal ridges: A pair of low, short to long, narrow ridges onthe midline of the caudal peduncle extending anteriorly fromthe anal fin base.Precaudal fins: All fins in front of the caudal fin.Precaudal pit: A depression at the upper and sometimeslower origin of the caudal fin where it joins the caudalpeduncle.Precaudal tail: That part of the tail from its base at the ventto the origins of the caudal fin.Precaudal vertebrae: Vertebrae from the occiput to thedorsal origin of the caudal fin.Predorsal ridge: A low narrow ridge of skin on the midline ofthe back anterior to the first dorsal fin base.Preorbital canals: On the neurocranium, anterior passagesfor the superficial opthalmic nerves out of the orbits and ontothe nasal capsules and rostrum, situated at the anteromesialedges of the supraorbital crests at the rear bases of thepreorbital processes; sometimes greatly expandedposteriorly.Preorbital processes: On the neurocranium, anterolateralprojections of the supraorbital crests, below which thepreorbital walls originate.Preorbital walls: On the neurocranium, the anteriorboundaries of the orbits, curved vertical plates of cartilagethat vary from complete to absent in neoselachians.Preorbitalis muscles: Paired head muscles that primitivelyoriginate on the rear of the nasal capsules or on thepreorbital walls, run diagonally rearwards, and insert on theadductor mandibulae at the mouth angles. Orectoloboidsand heterodontoids have the preorbitalis vertical, withcross-biased fibres in the latter, and the insertions are alongthe ventral edge of Meckel’s cartilage. In derivedorectoloboids the origins of the preorbitalis are expandedonto the cranial roof and the muscles greatly expanded.

26 FAO Species Catalogue for Fishery Purposes No. 1Primitively the preorbitalis may primarily serve to protrudethe jaws, but they may primarily serve to increase the powerof the bite in orectoloboids and heterodontoids. Also termedlevator labii superioris muscles.Preventral margin: In the caudal fin, the margin from thelower origin to the ventral tip of the caudal fin.Pristiophoroid: A saw shark, order Pristiophoriformes,family Pristiophoridae.Propterygium: In the pectoral fin skeleton of livingneoselachians, the anteriormost basal cartilage, adjacent tothe anterior edge of the mesopterygium and with one ormore radials attached to its distal end. In batoids withexpanded anterior pectoral fin lobes it becomes expandedand segmented into a propterygial basal and propterygialaxis, similar to the metapterygial basal and axis.Proximal: In any direction, at the near end of a structure.Pseudopera: On the external clasper glans, a dorsallyopening blind pocket along the lateral edge of the clasper,and about opposite the anterior edge of the glans.Pseudosiphon: On the external clasper glans, a dorsallyopening blind pocket along the medial edge of the clasper,and about opposite the cover rhipidion.Pterotic horn or process: On the neurocranium, elongatedposterior projections of the sphenopterotic ridges of the oticcapsules.Puboischiadic bar: A transverse flattened or cylindricalplate in the posterior body wall opposite the anterior ends ofthe pelvic fins, in front of the vent and at the posterior end ofthe body cavity, that supports a few anterior pelvic radialsand a basal cartilage, the basipterygium. The pelvic girdle.Radial cartilages or radials: The small, segmented, moredistal cartilages of the precaudal fins, attached proximally tothe distal edges of the basal cartilages. In the pectoral finskeleton of living neoselachians, the radials mostly havethree segments but range from no segments to 30 or more.The radial segments adjacent to the pectoral basals are theproximal radials, the radial segments furthest from thebasals are the distal radials, and any segments betweenthem are intermediate radials.Radii: In a vertebral centrum in cross-section, branchingplates of calcified cartilage in the intermedialia. These havea radial orientation from the centre of the centrum.Ray: See batoid.Replacement series: A series of oral teeth that are lingualto the functional series, and not in a functional position onthe jaw.Rhipidion: In nonbatoid sharks, a longitudinal, elongatedflap attached to the floor of the glans along its base and withits free edge directed laterally. In skates (Rajoidei) rhipidionis used for a soft mass of erectile tissue in the glans, notnecessarily homologous to the rhipidion of nonbatoidsharks.Rhomboidal: In the form of a rhombus or diamond.Ribs: On the shark vertebral column, short to elongatedpaired and typically pointed cartilages attached to thebasiventral cartilages and extending into the horizontalseptum of the segmented trunk musculature or myomeres.Chondrichthyan ribs are therefore dorsal ribs rather thanventral ribs as in bony fishes (which support the bodycavity).Ring valve: A type of spiral intestinal valve in which thevalve turns are very numerous and short and resemble astack of washers.Root lobe: Sharks often have the roots of their oral teethdivided into separate lobes at their midlengths, which aretermed mesial and distal root lobes.Root: The proximal part of the oral tooth, made of porousosteodentine and anchoring the tooth in the dentalmembrane of the jaw.Rostral keel: In the neurocranium of squaloids, a largevertical plate on the underside of the rostrum and internasalseptum, sometimes reduced, and with the cavities of thesubnasal fenestrae on either side of the keel.Rostral node: On the neurocranium, the anterior end of therostrum of cartilaginous fishes, and the plate formed by thefused anterior ends of the tripodal rostra in manygaleomorph sharks.Rostromandibular muscle: In the orectoloboid familyParascylliidae, paired head muscles that originate on thesides of the adductor mandibulae muscles and insert vialong tendons on the medial rostral cartilage. These arepossibly for depressing the snout. Not found in any othersharks, though analogous muscles exist in batoids.Rostronuchal muscles: In the orectoloboid familyParascylliidae, paired head muscles that originate on thedorsal myomeres of the nape, and insert via long tendons onthe medial rostral cartilage. These are possibly for elevatingthe snout. Not found in any other sharks, though analogousmuscles exist in batoids.Rostrum: On the neurocranium, the cartilaginousanteriormost structure which supports the prenasal snoutincluding lateral line canals and masses of ampullae, and islocated in front of the nasal capsules and anterior fontanelle.The rostrum is very variable, and in squalomorph sharks isprimitively trough or basin-shaped, while it may beprimitively rod-shaped or tripodal in galeomorph sharks. It isabsent in a few nonbatoid sharks and in many batoids. Seerostrum, tripodal.Rostrum, tripodal: The rostrum of the neurocranium inlamnoid and carcharhinoids is primitively tripodal, with a pairof dorsolateral lateral rostral cartilages that arise from theposterolaterodorsal surfaces of the nasal capsules or fromthe preorbital wall, and a medial rostral cartilage thatarises from the anteromedial surface of the internasalseptum. The medial and lateral rostral cartilages extendanteriorly and articulate or fuse at the rostral node. Livingorectoloboids have only the medial rostral cartilage althougha tripodal rostrum may be present in some fossilorectoloboids, while heterodontoid sharks lack a rostrum asadults but apparently lose it as embryos.Row: In oral teeth, a single replicating line of teeth,approximately transverse to the longitudinal jaw axis, whichincludes functional teeth and their replacements, derivedfrom one tooth-producing area on the jaw.

Sharks of the World, Vol. 2 27Saw or saw-snout: The elongated snout in sawfish andsawsharks, with side and (in sawsharks) ventral teethformed from enlarged denticles, used to kill, ensnare or digfor prey. Also termed rostral saw.Scapulocoracoid: The primitively U-shaped cartilage in thebody wall just behind the gills and at the anterior end of thepectoral bases, that supports the pectoral fins andarticulates with the pectoral basals. The scapulocoracoidconsists of a ventral coracoid bar connecting its pairedlateral faces with articular condyles or ridges for thepectoral basals, and a pair of dorsal scapular processesdorsal to the lateral faces. The scapular processessometimes have separate suprascapulae above them, butthey are sometimes fused with the scapular processes. Thecoracoid bar has a medial joint or even a separate medialcartilage (sternal cartilage) in a few living sharks, as withmany fossil cartilaginous fishes. The pectoral or shouldergirdle.Scroll valve: A type of spiral intestinal valve in requiem andhammerhead sharks in which the valve has uncoiled andresembles a rolled-up bib or scroll.Second dorsal fin: The posteriormost dorsal fin of two incartilaginous fishes, ranging in position from over the pelvicfinbases to far posterior on the precaudal tail.Secondary caudal keels: Low horizontal dermal keels onthe ventral base of the caudal fin in mackerel sharks(Lamnidae) and sometimes somniosids.Secondary lower eyelid: The eyelid below or lateral to thenictitating lower eyelid, separated from it by a suboculargroove or pocket, and, in many carcharhinoids with internalnictitating lower eyelids, functionally replacing them as lowereyelids. Some orectoloboids have shallow suboculargrooves separating their non-nictitating lower eyelids fromweakly developed secondary lower eyelids. They may,however, be able to close their eye openings by retractingthe eyeballs.Semiplesodic fin: In some sharks, a pectoral or dorsal finwith the fin radial cartilages extending partway into the finweb but not to its distal edges, essentially intermediatebetween plesodic and aplesodic fins.Series: In oral teeth, a line of teeth along the jaws which isparallel to the jaw axis and includes teeth from all rowspresent.Serrations: In oral teeth, minute teeth formed by the cuttingedge of the crown that enhance the slicing abilities of theteeth.Shark: Generally used for cylindrical or flattenedcartilaginous fishes with 5 to 7 external gill openings on thesides of their heads, pectoral fins that are not attached to thehead above the gill openings, and a large, stout tail with alarge caudal fin; that is, all living elasmobranchs except therays or batoids. Living sharks in this sense are all membersof the Neoselachii, the modern sharks and rays. Shark isalso used loosely for fossil chondrichthyans that are notneoselachians but have a shark-like form, and even for‘spiny sharks’ (acanthodians) and for certain teleosts. Raysare essentially flattened sharks with the pectoral finsattached to their heads and are cladistically nested withinthe squalomorph sharks, while living chimaeras are theimmediate sister group of living neoselachians and arecalled ghost sharks or silver sharks. Hence shark is usedhere in an alternate and broader sense to include the raysand chimaeras.Shoulder: In oral teeth, an arcuate or straight,convex-edged section of the crown foot, without cuspletsand similar to a blade but without a cutting edge.Single oviparity: A mode of egg-laying or oviparity in whichfemale sharks produce encased eggs in pairs, which are notretained in the oviducts and are deposited on the bottom.Embryos in the egg-cases are at an early developmentalstage, and take a few months to over a year to hatch. Foundin almost all oviparous cartilaginous fishes.Siphons: A pair of dermal sacs in the ventral abdominal wallof male sharks, connecting posteriorly with the apopyles ofthe claspers, and extending anteriorly a variable distancefrom about opposite the pelvic origins to opposite thepectoral bases.Skull or cranium: The skull or head skeleton of sharksincludes the neurocranium and the splanchnocranium orvisceral arches. The visceral arches articulate with and areassociated with the neurocranium, but, except for the upperjaws of many holocephalans, are not fused to it. Also termedsyncranium.Snout: That part of a cartilaginous fish in front of its eyesand mouth, and including the nostrils.Sphenopterotic ridge: On the neurocranium, a horizontalridge along the dorsolateral edge of each otic capsule thateither ends at the occiput or terminates in an expandedpterotic process.Spiracle: A small to large opening between the eye and firstgill opening of most sharks and rays, representing themodified gill opening between the jaws and hyoid (tongue)arch. This is secondarily lost in chimaeras and some sharks.Spiral or conicospiral valve: An intestinal valve shapedlike a corkscrew or augur, with the valve angled anteriorlyand medially in the intestine.Splanchnocranium: That part of the shark skull includingthe visceral arches. These include the jaws or mandibulararch, the tongue or hyoid arch, and the five to seven gill orbranchial arches. Also, viscerocranium.Squalene: A long-chain oily hydrocarbon present in the liveroil of deepwater cartilaginous fishes. It is highly valued forindustrial and medicinal use.Squaloid: A dogfish shark, a member of the orderSqualiformes, including bramble sharks, spiny dogfish,gulper sharks, lantern sharks, viper sharks, rough sharks,sleeper sharks, kitefin sharks, and cookiecutter sharks.Squalomorph: Referring to the Squalomorphii.Squalomorphii: The neoselachian superorderSqualomorphii, including the hexanchoid, squaloid,squatinoid, pristiophoroid, and batoid sharks.Squatinoid: An angel shark, order Squatiniformes, familySquatinidae.Squatinoid caudal fin: Angel sharks (Squatiniformes) areunique among living sharks in having hypocercal caudal fins

28 FAO Species Catalogue for Fishery Purposes No. 1that resemble inverted caudal fins of ordinary sharks. Thedorsal margin is subdivided into a predorsal margin fromthe upper origin to its dorsal tip (analogous to the preventralmargin and ventral tips in ordinary sharks), a postdorsalmargin (like the postventral margin) from the dorsal tip to itssupraterminal notch (similar to the subterminal notch), and ashort supraterminal margin and large ventral terminalmargin (similar to the subterminal and terminal margins)between the supraterminal notch and the ventral tip of thecaudal. The ventral margin has a preventral margin forminga ventral lobe with the ventral tip and the ventral terminalmargin.Stapedial foramen or fenestra: On the neurocranium, aforamen through the posteroventromedial surface of eachsuborbital shelf into the orbit, for the stapedial or orbitalarteries. It may be greatly expanded into a stapedial fenestrain sharks with greatly coiled stapedial arteries or lost insharks with the suborbital shelves greatly reduced orabsent.Stapediocarotid foramen: On the neurocranium of certainsharks, fusion of the stapedial and carotid foramina on eitherside produces a single pair of stapediocarotid foramina.Subcaudal keel: In a few dogfish sharks (familyCentrophoridae), a single longitudinal dermal keel on theunderside of the caudal peduncle.Subethmoid fossa: On the neurocranium, a deep cavity onthe ventral surfaces of the nasal capsules and the internasalplate, into which fit the palatine processes of the upper jaws.Subnasal fenestrae: On the neurocranium of squaloids, apair of apertures in the internasal plate between the nasalcapsules that connect the cerebral cavity with two ventralfluid-filled cavities between the nasal capsules and therostral keel. The fenestrae themselves are covered by toughmembranes as with the anterior fontanelle. Subnasalfenestrae are present in most squaloids but reduced in a fewderived species, and are not found in other sharks. Theirfunction is obscure but may be sensory. Also termed basalcommunicating canals.Suborbital shelf: On the neurocranium, a horizontal platearising on the ventral junction of the orbital wall and basalplate on each side which extends from the nasal capsule tothe otic capsule; it forms the floor of the orbit. Awell-developed suborbital shelf is apparently primitive forshark-like fishes but is variably telescoped, reduced or lostin many squalomorph sharks and a few galeomorphs.Subterminal margin: In the caudal fin, the margin from thesubterminal notch to the ventral beginning of the terminalmargin.Subterminal mouth or ventral mouth: Mouth located onthe underside of the head, behind the snout. Also termed aninferior mouth, in reference to its ventral position but not itsfunction. A superior mouth (not found in living cartilaginousfishes) is on the dorsal surface of the head.Subterminal notch: On the caudal fin of most nonbatoidsharks and at least one batoid, the notch in the lower distalend of the caudal fin, between the postventral andsubterminal margins, and defining the anterior end of theterminal lobe.Superficial ophthalmic nerve foramina: Foramina for theroots of the superficial ophthalmic nerves in the medial wallof the orbits, separate from the orbital fissure. Theseforamina are confluent with the orbital fissure in manysharks.Supraorbital crest: On the neurocranium, an archedhorizontal plate of cartilage forming the dorsal edge of theorbit on each side; it arises from the medial orbital wall andthe cranial roof and extends horizontally from the preorbitalprocess to the postorbital process. It is apparently primitivefor shark-like fishes but is variably reduced or absent insome living elasmobranchs.Supraorbital or brow ridge: A dermal ridge above eacheye, particularly well-developed in heterodontoids and someorectoloboids.Symphyseal or symphysial groove: A longitudinal grooveon the ventral surface of the lower jaw of some orectoloboidsharks, extending posteriorly from the lower symphysis.Symphysial teeth: Larger oral teeth in one row on eitherside of the symphysis, distal to medials or alternates wherepresent. Symphysials are broader than medials and usuallyhave asymmetrical roots.Symphysis: The midline of the upper and lower jaws, wherethe paired jaw cartilages articulate with each other.Syntype: Two or more specimens used and mentioned inan original description of a species, where there was nodesignation of a holotype or a holotype and paratype(s) bythe describer of the species.Systematics: Scientific study of the kinds and diversity oforganisms, including relationships between them.Tail: That part of a cartilaginous fish from the cloacalopening or vent (anus in chimaeroids, which lack a cloaca)to the tip of the caudal fin or caudal filament, and includingthe anal fin, usually the second dorsal fin when present, andcaudal fin.Taxon, plural taxa: A taxonomic group at any level in aclassification. Thus the taxon Chondrichthyes is a class withtwo taxa as subclasses, Elasmobranchii and Holocephali,and the taxon Galeorhinus, a genus, has one taxon as aspecies, G. galeus.Taxonomy: Often used as a synonym of systematics orclassification, but narrowed by some researchers to thetheoretical study of the principles of classification.Term foetus: See foetus.Terminal 3 cartilage: A wedge-shaped or elongatedcartilage articulating with the posterior edge of the ventralmarginal cartilage and over the ventral terminal cartilages. Itsupports a variety of structures, including clasper spinesand spurs, the shields of many skates (Rajoidei), and themesorhipidion of some carcharhinoid sharks.Terminal lobe: In the caudal fin of most nonbatoid sharksand at least one batoid, the free rear wedge-shaped lobe atthe tip of the caudal fin, extending from the subterminalnotch to the posterior tip.

Sharks of the World, Vol. 2 29Terminal margin: In the caudal fin, the margin from theventral end of the subterminal margin to the posterior tip.Terminal mouth: Mouth located at the very front of theanimal. Most cartilaginous fishes have subterminal mouths,but some species (viper sharks, wobbegongs, angel sharks,frilled sharks, whale sharks, megamouth sharks, andManta) have it terminal or nearly so.Thorn: In many batoids, most angel sharks and the brambleshark (Echinorhinus brucus), enlarged, flat conicaldenticles with a sharp, erect crown and a flattened base(which may grow as the shark grows).Tongue arch: See hyoid arch.Transverse groove: In oral teeth, a deep groove transverseon the lingual root surface, transecting it into mesial anddistal root lobes.Transverse notch: In oral teeth, a distinct notch in theproximal labial edge of the root at about its midlength.Transverse ridges: Small narrow ridges on the labial andlingual surfaces of the crown, apicobasally oriented andsometimes extending to the cusp edges.Tribasal pectoral fin: A pectoral fin skeleton with threebasal cartilages, the propterygium, mesopterygium, andmetapterygium, primitively found in most euselachiansincluding living neoselachians.Trilobate lower lip: In advanced orectoloboids, shalloworolabial grooves divide the lower lips into a medial sectionand a pair of lateral sections.Tropeic folds: Longitudinal paired ridges on the ventralmidline of the abdomen in frilled sharks(Chlamydoselachidae).Truncate: Blunt, abbreviated.Trunk: That part of a cartilaginous fish between its head andtail, from the last gill openings to the vent, including theabdomen, back, pectoral and pelvic fins, and often the firstdorsal fin.Umbilical cord: A modified yolk stalk in placental viviparoussharks, carrying nutrients from the placenta to the foetus.Unpaired fins: The dorsal, anal, and caudal fins.Upper eyelid: The dorsal half of the eyelid, separated by adeep pocket (conjunctival fornix) from the eyeball. Theupper eyelid fuses with the eyeball and the pocket is lost inall batoids.Upper origin: In the caudal fin, the anterodorsal beginningof the epaxial or upper web of the caudal fin, at the posteriorend of the dorso-caudal space (see measurementillustrations).Upper postventral margin: In the caudal fin, the upper partof the postventral margin of the hypaxial web, from theposterior notch to the subterminal notch.Uterine cannibalism or cannibal viviparity: A mode ofreproduction in which foetuses deplete their yolk-sacks earlyand subsist by eating nutritive eggs produced by the mother(see oophagy) or first eat smaller siblings and then nutritiveeggs (see adelphophagy).Vent: The opening of the cloaca on the ventral surface of thebody between the inner margins and at the level of the pelvicfin insertions.Ventral: Downward, in the vertical direction of the abdomen.See dorsal.Ventral fin: See pelvic fin.Ventral lobe: In the caudal fin, the expanded distal end ofthe preventral and lower postventral margins, defined by theposterior notch of the caudal fin.Ventral margin: In the caudal fin, the entire ventral marginfrom lower origin to posterior tip, either a continuous marginor variably subdivided into preventral, postventral,subterminal and terminal margins.Ventral marginal: In the clasper skeleton, a flatsemicylindrical cartilage that is partially fused to the lateraledge of the axial cartilage, and forms the lateral wall of theclasper groove.Ventral terminal: On the skeleton of the clasper glans, anoften triangular, elongated, curved, plate-like cartilage thatarticulates or is attached to the lateral or ventrolateral edgeof the end-style and to the posterior end of the ventralmarginal cartilage.Ventral tip: In the caudal fin, the ventral apex of the caudalfin where the preventral and postventral margins merge.Vertebra, plural vertebrae: A single unit of the vertebralcolumn, including a vertebral centrum and associatedcartilages that form neural arches and ribs or haemalarches.Vertebral axis: That part of the vertebral column inside thebase of the caudal fin.Vertebral column: The entire set or string of vertebrae or‘backbone’ of a shark, from the rear of the chondrocraniumto the end of the caudal base. Living elasmobranchs rangefrom having as few as 60 vertebrae (some squaloids of thefamily Dalatiidae) to as many as 477 vertebrae (threshersharks).Visceral arches: See splanchnocranium.Viviparity: Used in two ways in recent literature, as beingequivalent to placental viviparity only, that is forcarcharhinoid sharks with a yolk-sac placenta; or for allforms of live-bearing or aplacental viviparity.Web, fin: See fin web.Yolk sac or yolk sack: Almost all sharks start embryonicdevelopment somewhat like a chicken, as a large sphericalyolky egg inside an elongated shell, the egg case. A smalldisk of dividing cells represents the pre-embryo or blastulaatop the huge yolk mass. The blastula expands around thesides and ventral surface of the yolk mass, and differentiatesinto an increasingly shark-like embryo, the yolk sac orbag-like structure containing the yolk, and a narrow tubularyolk stalk, between the abdomen of the embryo and the yolksac.

30 FAO Species Catalogue for Fishery Purposes No. 1Yolk stalk: The connecting passage between embryo orfoetus and yolk sac, which allows yolk to pass from the sacinto the embryonic gut.Yolk-sac placenta: An organ in the uterus of some groundsharks (order Carcharhiniformes), formed from theembryonic yolk-sac of the embryo and maternal uterinelining, through which maternal nutriment is passed to theembryo. It is analogous to the placenta of live-bearingmammals. There are several forms of yolk-sac placentas incarcharhinoid sharks, including entire, discoidal, globular,and columnar placentas (see Compagno, 1988).Yolk-sac viviparity: Live-bearing in which the young arenourished primarily by the yolk in the yolk sacs, which isgradually depleted and the yolk sacs reabsorbed until theyoung are ready to be born.click for next page

click for previous pageSharks of the World, Vol. 2 312. SYSTEMATIC CATALOGUE2.1 Order HETERODONTIFORMES - Bullhead sharksOrder: [Group] Heterodonti Garman, 1885, Bull. Mus. Comp. Zool. Harvard, 12(1): 30, emended to OrderHeterodontiformes.Number of Recognized Families: 1.Synonyms: [Part] 1 Squali, Abtheilung [Division] 2: Müller and Henle, 1838: 27; Müller and Henle, 1839: 27; also [Part] 1Squali, Abtheilung [Division] 2, Unterabtheilung [Subdivision] 3: Müller and Henle, 1839: 66. Ordo Plagiostomi, SubordoSqualini, Sectio Proktopterides, Tribus Dinotopterini: Bleeker, 1859: xi. Order Squali, Suborder Squali: Gill, 1862b: 396.Order Squali, Suborder Galei: Gill, 1872: 23. Order Plagiostomi diplospondyli, Suborder Plagiostomi Asterospondyli, Group3 Acrodonten: Hasse, 1879: 50. Order Selachii, Suborder Asterospondyli: Woodward, 1889: 157. Order Prosarthri: Gill,1893: 129. Order Asterospondyli, Suborder Proarthri: Jordan and Evermann, 1896: 19. Order Euselachii, SuborderPleurotremata, Division Squaloidei: Regan, 1906a: 723. Order Selachii, Group 2, Division A, Suborder Heterodonti:Goodrich, 1909 (in part): 143. Order Plagiostoma, Suborder Antacea, “Group” Centracoidei: Garman, 1913: 11, 13. OrderPleurotremata, Suborder Squaloidei: Engelhardt, 1913: 100. Order Centraciones, Suborder Prosarthri: Jordan, 1923: 95.Order Heterodontea, Suborder Heterodontida, Superfamily Heterodontoidea: White, 1936: 4; White, 1937: 37, tab. 1;Whitley, 1940: 69. Order Euselachii, Suborder Heterodontiformes: Bertin 1939a: 9. Order Heterodontiformes: Berg, 1940:134, 135; Berg and Svetovidov, 1955: 61; Arambourg and Bertin, 1958: 2028; Patterson, 1967: 667; Lindberg, 1971: 8, 256;Rass and Lindberg, 1971: 303; Compagno, 1973: 28; Applegate, 1974: 743; Nelson, 1976: 32; Chu and Meng, 1979: 114,tab. 2; Compagno, 1984: 154; Nelson, 1984: 50; Gubanov, Kondyurin and Myagkov, 1986: 3, 42; Cappetta, 1987: 26, 69;Compagno, 1988: 382; Nelson, 1994: 45; Shirai, 1996: 32; Eschmeyer, 1998. Order Heterodontiformes, SuborderHeterodontoidei: Berg, 1940: 134, 135; Berg and Svetovidov, 1955: 61. Order Asterospondyli: Fowler, 1941: 4, 13; Smith,1949: 37, 39. Order Selachii, Suborder Heterodontoidea: Romer, 1945: 576; Bigelow and Schroeder, 1948: 95; Romer,1966: 349. Order Heterodontoidea: Schultz and Stern, 1948: 224. Order Pleurotremata, Suborder Heterodontiformes:Budker and Whitehead, 1971: 5, tab. 2. Order Heterodontida, Suborder Heterodontina, Superfamily Heterodonticae:Fowler, 1966: 321, 330, 331. Order Pleurotremata, Suborder Squaloidea: Norman, 1966: 24. Order Hexanchida, SuborderHeterodontoidei: Glikman, 1967: 214. Order Selachii: Blot, 1969: 702-776. Order Heterodontiformes, SuborderHeterodontoidea: Chu and Meng, 1979: 114, tab. 2. Order Galeomorpha, Suborder Heterodontoidea: Carroll, 1988: 598.FAO Names: En - Bullhead sharks.Field Marks: The only living sharks with two spined dorsal fins and an anal fin; also a pig-like snout, small anterior mouth,enlarged molariform teeth in the back of the mouth, supraorbital ridges, rough skin, paddle-like paired fins, and enlarged firstgill slits.Diagnostic Features: Head elevated with crests above eyes, not depressed or expanded laterally. Snout very short, slightlydepressed and bluntly rounded, and without lateral teeth or rostral barbels. Eyes dorsolateral on head, without nictitatinglower eyelids, secondary lower eyelids, or subocular pouches; upper eyelids not fused to eyeball. Spiracles very small, justbehind or about opposite eyes but well below eye level. Five pairs of gill openings present on sides of head, withposteriormost two or three behind pectoral-fin origins. Nostrils diagonal on front of snout, without barbels but with prominentcircumnarial grooves around incurrent apertures; nostrils with deep nasoral grooves connecting excurrent apertures tomouth, anterior nasal flaps moderately long and reaching mouth. Mouth small, almost terminal on head, broadly arched andshort, ending in front of posterior corners of eyes. Labial furrows large and present on both jaws. Teeth strongly differentiatedalong the jaws, with small anterolateral teeth and enlarged molariform posterior teeth; no gap or small intermediate teethbetween anterior and lateral teeth in the upper jaw; anterolateral teeth with orthodont histological structure but posteriorteeth osteodont. Trunk cylindrical, not flattened and ray-like. Caudal peduncle without lateral dermal ridges or keels. Dermaldenticles covering entire body, not enlarged as thorns or spines. Pectoral fins moderately large, not expanded and ray-like,without triangular anterior lobes that cover the gill openings. Pectoral girdle (scapulocoracoid) high, U-shaped, without amedial joint, and with superscapulae directed dorsally and not contacting vertebral column. Pectoral-fin skeleton primitivelytribasal (dibasal with propterygium fused to mesopterygium in some species), with propterygium in contact with radials andmetapterygium without a proximal segment; pectoral fins semiplesodic, with radials extending into the basal fin webs; radialcount 15 to 19 with mostly 2 to 8 segments. Pelvic fins small, with vent continuous with their inner margins. Claspers withshort siphons in the abdomen at the pelvic-fin bases but without clasper sacs; clasper glans with a large pseudosiphon, acover rhipidion, a rhipidion and clasper spine; dorsal and ventral marginals of clasper skeleton well-developed but with onlythe dorsal marginal partially rolled into a tube for clasper canal. Two spine-bearing dorsal fins present, with origin of first wellin front of pelvic-fin bases and over pectoral fins; dorsal-fin skeleton with segmented radials articulating with enlargedintermediate cartilages and a single basal plate that also carries the large fin spine. Anal fin present. Caudal fin with a longdorsal lobe and a short ventral lobe; vertebral axis elevated into the dorsal caudal lobe (heterocercal caudal fin). Vertebralcalcification strong, secondary calcification in form of several radii that extend under the basals but without intermedialwedges, annuli, or diagonal calcifications. Total vertebral count 103 to 123, precaudal vertebrae 60 to 81. Neurocraniumwithout a rostrum; nasal capsules trumpet-shaped and without subnasal fenestrae (basal communicating canals) or

32 FAO Species Catalogue for Fishery Purposes No. 1antorbital cartilages; orbits with incomplete preorbital walls, strong supraorbital crests, strong suborbital shelves, separateforamina for hyomandibular nerves but no separate foramina for superficial ophthalmic nerves, and incomplete postorbitalwalls without lateral commissures for lateral head vein; occipital condyles low but broad, with a prominent occipitalhemicentrum between them. Jaws elongated, upper jaws (palatoquadrates) with low, ridge-like orbital processes thatarticulate with nasal capsules and orbits in horizontal grooves contacting ethmoid region, basal plate and suborbital shelves;orbital processes not penetrating supraorbital crests. Hyobranchial skeleton with moderately broad, elongated basihyoid;posterior two pharyngobranchials and last epibranchial fused as a yoke-shaped element. Head muscles include broadenlarged vertical preorbitalis with cross-biased fibres; narrow levator palatoquadrati and separate first dorsal constrictorunder postorbital process and not extending behind orbits; adductor mandibulae muscles not segmented and not notchedanteriorly for mouth gape; a discrete craniomandibular muscle between the lower jaw and orbital walls; nomandibulocutaneous muscle between upper jaw and skin; and no postocular eyelid muscles. Intestinal valve of conicospiraltype, with seven turns. Reproduction oviparous (egg laying), with unique screw-shaped egg cases.Distribution, Habitat, Biology, Interest to Fisheries and Human Impact, and Local Names: See family Heterodontidaebelow.Remarks: The Heterodontiformes are a small but highly distinctive group of sharks that have generally been recognized asa discrete taxon at the genus, family, and higher levels. They have a long fossil record extending to almost the beginning ofthe Mesozoic era and agree with hybodont sharks and other extinct euselachians in having stout dorsal-fin spines and ananal fin.Bullhead sharks were first discovered in Australia in the late eighteenth century in the form of the Port Jackson shark(Squalus portus jacksoni Meyer, 1793 and several synonyms). This species was initially classified in the Linnaean sharkgenus Squalus but soon was accorded two genus-group names, Heterodontus Blainville, 1816 and its junior synonymCestracion Oken, 1817, which initially were proposed as subgenera of Squalus but were recognized as genera by mostsubsequent authors. Bonaparte (1838) proposed the new subfamily Cestraciontini (family Squalidae) and Müller and Henle(1839) proposed a new family Cestraciontes for Cestracion, while Gray (1851) proposed the new tribe Heterodontina forHeterodontus which was raised in rank to the family Heterodontoidae by Gill (1862b).Many nineteenth and early twentieth century writers used the genus Cestracion in favour of Heterodontus and recognizedthe family Cestracionidae (or variants), including Müller (1845, family Cestraciontes), Bleeker (1859, familyCestracionoidei), Dumeril (1865, family Cestraciontes), Owen (1866), Günther (1870), Woodward (1889, 1898), Zittel et al.(1902), Regan (1906a), Goodrich (1909), Garman (1913, family Centraciontidae), Engelhardt (1913), and Zittel, Broili andSchlosser (1923). Swainson (1839) placed Cestracion in the family Squalidae, subfamily Centrininae along with varioussqualoids and other sharks. Van der Hoeven (1858) placed Cestracion in the family Selachii along with all other sharks whileHasse (1879) placed it in his Group 3, Acrodonten without familial assignment. The use of Heterodontus as a seniorsynonym of Cestracion began in the second half of the nineteenth century and became universal for most of the twentiethcentury. The family Heterodontidae was recognized by Gill (1872, 1893), Jordan and Gilbert (1883), Jordan and Evermann(1896), Bridge (1910), Jordan (1923), White (1936, 1937), Bertin (1939a), Berg (1940), Whitley (1940), Fowler (1941, 1966),Romer (1945, 1966), Bigelow and Schroeder (1948), Schultz and Stern (1948), Smith (1949), Berg and Svedovidov (1955),Matsubara (1955), Arambourg and Bertin (1958), Norman (1966), Glickman (1967), Patterson (1967), Blot (1969), Bailey etal. (1970), Rass and Lindberg (1971), Budker and Whitehead (1971), Pinchuk (1972), Compagno (1973, 1981b, 1984),Nelson (1976, 1984, 1994), Chu and Meng (1979), McEachran and Compagno (1982), Gubanov, Kondyurin and Myagkov(1986), Cappetta (1987), Carroll (1988), Eschmeyer (1990, 1998), and Shirai (1992, 1996).The Heterodontidae (or Cestraciontidae) were often grouped with various fossil shark groups by earlier authors, particularlythe hybodonts, and with various families of living sharks including lamnoids, carcharhinoids, and squaloids. The concept of aseparate ordinal-group for Heterodontidae dates from Garman (1885) as a “group” Heterodonti, while Gill (1893) included itin a separate order Prosarthri (suborder Proarthri of Jordan and Evermann, 1896 and of Jordan, 1923, who also recognizedan order Centraciones). Goodrich (1909) included Heterodontidae in the suborder Heterodonti, and Garman (1913)included it in the “group” Centracoidei. White (1936, 1937) recognized the order Heterodontea, which was listed by Bertin(1939a) as the suborder Heterodontiformes, by Berg (1940) as the order Heterodontiformes, and by Bigelow and Schroeder(1948) as the suborder Heterodontoidea. Most subsequent writers tend to accord it a separate order Heterodontiformes,including the writer (Compagno 1973, 1977, 1984, 1988, 1999, and present work).The anatomy of heterodontoids was described by Gegenbaur (1865, 1872), Haswell (1885), Daniel (1914, 1915, 1928),Holmgren (1941), Kesteven (1942), Compagno (1973, 1977), Shirai (1992, 1996), and de Carvalho (1996), while generalmorphology and systematics were summarized by Dumeril (1865), Miklouho-Maclay and Macleay (1879, 1886), Günther(1870), Regan (1908b), Garman (1913), Smith (1942), Taylor (1972), and Compagno (1984). It is likely that theheterodontoids are related to the lamnoids, carcharhinoids and orectoloboids, with the most likely hypothesis being that theheterodontoids are a sister group to the three ‘galeoid’ orders rather than to the orectoloboids alone (see discussions inCompagno, 1973, 1977, 1988; Shirai, 1996; and de Carvalho 1996).

Sharks of the World, Vol. 2 332.1.1 Family HETERODONTIDAEFamily: Tribe Heterodontina Gray, 1851, List Fish British Mus., Pt. 1, Chondropterygii, British Mus. (Nat. Hist.): 65 (FamilySqualidae).Type Genus: Heterodontus Blainville, 1816. Raised to the rank of Family Heterodontoidae by Gill, 1862b, Ann. LyceumNat. Hist. New York, 7(32): 403.Number of Recognized Genera: 1.Synonyms: Subfamily Cestraciontini Bonaparte, 1838: 211 (Family Squalidae). Family Cestraciontes Müller and Henle,1839: 76. Type genus: Cestracion Cuvier = Oken, 1817. Family Centraciontidae Garman, 1913: 13, 180. Type genus:Centracion Gray, 1831, possible error for Cestracion Cuvier, = Oken, 1817. Family Heterondontidae Bass, D’Aubrey andKistnasamy, 1975d: 17. Apparent error for Heterodontidae, repeated twice on the same page but correctly spelledelsewhere (title page and table of contents, p. 2).FAO Names: En - Bullhead sharks, Horn sharks; Fr - Requins dormeurs; Sp - Dormilones.Field Marks: See order Heterodontiformes above.Diagnostic Features: See order Heterodontiformes above.Distribution: Bullhead sharks occur in warm-temperate and tropical continental waters of the western Indian Ocean,western and eastern Pacific, but are absent from the Atlantic and from oceanic insular waters.Habitat: These are warm-temperate and tropical bottom sharks of water above 21°C, mostly confined to the continental andinsular shelves and uppermost slopes. They occur from the intertidal to 275 m depth, but most are found in water shallowerthan 100 m.Biology: As far as is known, these are sluggish, rare to uncommon night-active sharks, slowly swimming and crawling onrocky, kelp-covered and sandy bottom. Some species at least favour rocky crevices and caves, where they spend the dayresting. At least one species is migratory in coastal waters when adult, and returns to its breeding sites each year after longmigrations.These sharks are oviparous, producing eggs in unique, large, spiral-flanged egg cases. At least two species lay eggs inspecific ‘nesting’ sites. Eggs may take over five months to hatch, and young hatch at a large size, over 14 cm.Bullhead sharks primarily feed on benthic invertebrates. Sea urchins (echinoids) are a favourite food, but crabs, shrimp andother crustaceans, abalone, top shells (Trochidae; Gastropoda) and other marine gastropods, oysters, polychaetes,sipunculid worms and more rarely small fish are also eaten.Interest to Fisheries and Human Impact: These sharks are of minimal interest to fisheries, being caught as a bycatch ofbottom trawl and line fisheries and utilized for human consumption and for fishmeal or discarded. They are commonlycaught by divers and sportsfishers and at least one species is prized by divers for its fin spines, which are made intojewellery.Several species of bullhead sharks have been kept in aquaria, where they have proved to be extremely hardy and can livefor over a decade. Mating, egg-laying, hatching of eggs, and growth to maturity can occur in captivity.Some bullhead sharks are encountered by divers, who have commonly harassed them. Although regarded as harmless,these sharks can and do snap when provoked and occasionally pursue and bite their tormentors. One species is the subjectof ecotouristic diving in California at present.At least two species of bullhead sharks may be declining as a result of fishing pressure. Most bullhead sharks are caught aslow-level bycatch of fisheries for abundant commercial species (such as shrimp). Their presence in tropical coastal waterswith heavy fishing pressure and habitat destruction (such as dynamite and poison-fishing of coral reefs), makes for possibleconservation problems in some areas where they occur, such as the Indo-West Pacific and the tropical eastern Pacific.Local Names: Bullhead sharks, Port Jackson bull-head sharks, Port Jackson sharks, Horn sharks (English); Requin de PortJackson (French); Husha k’o, Bulkophaaie (South Africa), Nekozame-ka (Japan); Akula rogataia, Rogatye akuly, Bych’iakuli (Russia); Tubarões dorminhocos (Mozambique).Remarks: Most authors have recognized a single genus, Heterodontus or its synonym Cestracion, but Gyropleurodus hasbeen used as a separate genus by some authors for eastern Pacific species (Heterodontus francisci and H. quoyi) orspecies with carinate teeth, following Gill (1863). Whitley (1931, 1940) separated the distinctive crested bullhead shark(H. galeatus), in its own genus Molochophrys, while Fowler (1934) coined a subgenus, Wuia, for the zebra bullhead shark(Heterodontus zebra). The systematic arrangement of the family Heterodontidae with a single genus Heterodontus followsTaylor (1972) and Compagno (1984). There is possible scope for subgeneric allocations within the family, but this shouldfollow an analysis of the phyletics of these sharks.Literature: Dumeril (1865); Günther (1870); Garman (1913); Fowler (1941, 1966); Smith (1942); Taylor (1972); Compagno(1984); Michael (1993); Last and Stevens (1994).

34 FAO Species Catalogue for Fishery Purposes No. 1Heterodontus Blainville, 1816Genus: Subgenus Heterodontus Blainville, 1816 (Genus Squalus Linnaeus, 1758), Bull. Sci. Soc. Philomat. Paris, (8): 121.Type Species: “Philippi”=Squalus philippi Bloch and Schneider, 1801, by monotypy, a junior synonym of Squalus portusjacksoni Meyer, 1793.Number of Recognized Species: 9.Synonyms: Genus Cestracion Oken, 1817: 1183. Latinization of “Les Cestracions. Cuv.” Cuvier, 1816: 129. Type species:Squalus philippi Bloch and Schneider, 1801, by monotypy? Genus Centracion Gray, 1831: 5. Type species: Centracionzebra Gray, 1831, by monotypy? Error or emendation of Cestracion Oken, 1817? Genus Cestralion Müller and Henle,1838: 85. Probable error for Cestracion Oken, 1817. Genus Heterodontes Gill, 1862b: 403. Obvious error for HeterodontusBlainville, 1816, as name is spelled correctly on same page. Genus Gyropleurodus Gill, 1863: 331. Type species:Cestracion francisci Girard, 1854, by monotypy (or original designation). Genus Tropidodus Gill, 1863: 489. Type species:Cestracion pantherinus Valenciennes, 1846, by original designation. Genus Molochophrys Whitley, 1931: 310. Typespecies: Cestracion galeatus Günther, 1870, by original designation. Subgenus Wuia Fowler, 1934: 233 (GenusHeterodontus Blainville, 1816). Type species: Centracion zebra Gray, 1831, by original designation. Genus TropidopusBeebe and Tee-Van, 1941: 118. Apparent error for Tropidodus Gill, 1863. Genus Cestraction Fowler, 1941: 17. Apparenterror for Cestracion Oken, 1817. Genus Cetracion Fowler, 1941: 17. Error for Cestracion Oken, 1817 or Centracion Gray,1831.Diagnostic Features: See family Heterodontidae above.Key to Species:1a. Supraorbital ridges very high (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . Heterodontus galeatus1b. Supraorbital ridges moderate to low (Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2supraorbital ridgesvery highsupraorbital ridgesmoderate to low1 st dorsal-fin origin behindpectoral-fin basesFig. 24 Heterodontus galeatusFig. 25 Heterodontus quoyi2a. First dorsal-fin origin behind pectoral-fin bases (Fig. 25) . . . . . . . . . . . . . . . . . . . . Heterodontus quoyi2b. First dorsal-fin origin over pectoral-fin bases (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33a. Body and fins spotted (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43b. Body and fins striped or banded (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 st dorsal-fin origin overpectoral-fin basesbody and fins spotteddark stripes or bandsFig. 26 Heterodontus francisciFig. 27 Heterodontus zebra

Sharks of the World, Vol. 2 354a. Body and fins with white spots in adultsand subadults (Fig. 28); hatchlings withthin curved parallel lines on body . . . .. . . . . . . . . . . . . . Heterodontus ramalheirawhite spots4b. Body and fins with dark spots and(particularly in young), darker saddles(Fig. 29) . . . . . . . . . . . . . . . . . . . . . 5Fig. 28 Heterodontus ramalheira5a. Back and sides with small dark spots lessthan a third of eye diameter (Fig. 26); nolight-coloured bar on interorbital surface ofhead . . . . . . . . . . . . Heterodontus francisci5b. Back and sides with larger dark spots ahalf eye diameter or more (Fig. 29); alight-coloured bar on interorbital surface ofhead . . . . . . . . . . . Heterodontus mexicanusdark spotsFig. 29 Heterodontus mexicanus6a. Body with a harness pattern of darkstripes (Fig. 30) . . . Heterodontus portusjacksoniharness pattern6b. Body with vertical dark bands orsaddles, not arranged in a harnesspattern (Fig. 27) . . . . . . . . . . . . . . . . . 77a. Background colour of dorsal surfacewhite or cream with a zebra pattern of22 to 36 narrow dark markings fromsnout to origin of caudal fin;anal-caudal space over twice anal-finbase (Fig. 27). . . . . . . . . . Heterodontus zebra7b. Background colour of dorsal surface tan tobrown with 5 to 14 broad or narrow,diffuse-edged markings from snout toorigin of caudal fin; anal-caudal space lessthan twice anal-fin base (Fig. 31). . . . . . . . . 8Fig. 30Heterodontus portusjacksoni11-14 dark markingsFig. 31 Heterodontus japonicus8a. Dorsal, pectoral and caudal fins withoutabruptly black tips or white apical spots;about 11 to 14 dark markings from snout toorigin of caudal fin, including broad darksaddles and narrow bands between them. . . . . . . . . . . . . . . Heterodontus japonicus4-5 broad dark saddleswhite apical spots8b. Dorsal and pectoral fins, and ventralcaudal-fin lobe, with abruptly black tips,dorsal fins with white apical spots; 4 or 5broad dark saddles from snout tip to originof caudal fin, without narrow dark bandsbetween them (Fig. 32) . . . . . . . . . .. . . . . . . . . . . . . Heterodontus sp. A (Oman)black tipsFig. 32 Heterodontus sp. A

36 FAO Species Catalogue for Fishery Purposes No. 1Heterodontus francisci (Girard, 1854) Fig. 33Cestracion francisci Girard, 1854, Proc. Acad. Nat. Sci. Philadelphia, 7(6): 196. Holotype: U.S. National Museum of NaturalHistory, apparently lost according to Taylor (1972, Rev. shark fam. Heterodontidae: 47). Type locality, Monterey Bay,California. Not listed in catalogue of USNM shark types by Howe and Springer (1993, Smiths. Contr. Zool., [540]: 1-19).Syntypes possibly USNM 933 (2) according to Eschmeyer (1998, Cat. Fish.: CD-ROM).Synonyms: Heterodontus californicus Herald, 1961: 49. Apparent error for H. francisci, which was cited correctly byHerald on p. 32.Other Combinations: Gyropleurodus francisci (Girard, 1854).FAO Names: En - Horn shark; Fr - Requin dormeur cornu; Sp - Dormilón cornudo.Fig. 33 Heterodontus francisciField Marks: Dorsal fins with spines, anal fin present, colour pattern of small dark spots less than one-third eye diameter onlight background, no light bar on interorbital space between supraorbital ridges, first dorsal-fin origin over pectoral-fin bases.Diagnostic Features: Supraorbital ridges moderately low, abruptly truncated posteriorly; interorbital space deeplyconcave, depth between ridges less than one-fourth eye length. Anterior holding teeth with a cusp and a pair of cusplets inadults, posterior molariform teeth strongly carinate and not greatly expanded and rounded. Pre-first dorsal-fin length 22 to27%, and anal-caudal space 4 to 8%, of total length. Lateral trunk denticles small and smooth, area behind first dorsal finwith about 200 denticles per cm 2in adults. Propterygium separate, not fused to mesopterygium. First dorsal-fin spinedirected obliquely posterodorsally in juveniles and adults; first dorsal-fin origin anterior to pectoral-fin insertions, over orslightly behind midbases of pectoral fins and well posterior to fifth gill openings; first dorsal-fin insertion well anterior topelvic-fin origin and well behind pectoral-fin insertion; first dorsal-fin free rear tip opposite or somewhat anterior to pelvic-finorigins; first dorsal fin moderately high and semifalcate in adults, height 9 to 14% of total length, slightly larger than pelvicfins. Second dorsal-fin origin over or slightly in front of pelvic-fin rear tips, second dorsal fin somewhat falcate and nearly aslarge as first dorsal fin. Anal fin subangular and weakly falcate, with apex reaching lower caudal-fin origin when laid back;anal-caudal space about equal to anal-fin base. Total vertebral count 103 to 123, precaudal count 65 to 76, monospondylousprecaudal count 30 to 38, diplospondylous precaudal count 32 to 46, pre-first dorsal-fin spine count 12 to 16, count fromdiplospondylous transition to second dorsal-fin spine 7 to 16. Egg cases with flat thin spiral flanges diagonal to case axis andno tendrils on case apices; flanges with five turns. A large species, mature between 59 and 122 cm. Colour: backgroundcolour of dorsal surface dark to light grey or brown with dark brown or black spots on body and fins, spots generally less thanone-third eye diameter; body without a dark harness pattern; head without a light bar on interorbital surface; small dark spotspresent below eye on a dusky patch; fins without abrupt dark tips and white dorsal-fin apices; hatchlings without whorls onfins and body, colour pattern as in adults although brighter.

Sharks of the World, Vol. 2 37Distribution: Warm-temperate and subtropical watersof the eastern Pacific: USA (Central and southernCalifornia), Mexico (Baja California, Gulf of California),and probably Ecuador and Peru. Off the USA it is mostcommon off southern California but ranges to MontereyBay and may occasionally penetrate as far north as SanFrancisco Bay (where it is not resident) during northerninfluxes of warm water.Habitat: A common benthic and epibenthic shark, foundon the eastern Pacific continental shelf most abundantlyat depths from 2 to 11 m but ranging from the intertidaldown to at least 150 m. Found on rocky bottomsincluding reefs, kelp beds, sandy draws between rocks,and on sand flats. On rocks it often occurs in deepcrevices and small caves, and ventures far into largeunderwater caverns. Juveniles shelter on sandy bottom,often near algae, rocks, detritus, or in feeding holesexcavated by bat rays (Myliobatis californica).Biology: The horn shark is sluggish, nocturnal, andmostly solitary, though small aggregations have beenseen by divers. It is seldom seen moving during thedaytime but commonly has its head in a crevice. Shortly after dusk this shark becomes active and apparently feeds mostly atnight, but ceases activity after dawn. Adults tend to return to the same resting place every day, but range at night over a smallhome range of roughly 0.1 hectare. According to Michael (1993) these sharks migrate into deeper water in winter, but it isuncertain if this occurs in the tropical part of their range. Experimentation with captive horn sharks indicates that their dielactivity pattern is controlled by light intensity. The broad, muscular paired fins of the horn shark are used as limbs forclambering on the bottom, and are highly mobile and flexible. Swimming is slow and sporadic.Courtship and copulation have been observed in captivity. The male horn shark chases the female until the latter is ready,then both drop to the bottom. The male grabs the female’s pectoral fin with his teeth and inserts a single clasper in hercloaca; copulation lasts 30 to 40 min. One to two weeks later eggs are laid by captive females, one of which laid two eggs perday at 11 to 14 day intervals for four months. In nature these sharks mate in December or January and females drop eggs inFebruary to April. Females normally deposit eggs under rocks or in crevices between them, but in captivity they drop eggs onthe bottom where the contents of egg cases may be subsequently sucked out and eaten by these sharks. Eggs can bereadily hatched in aquaria and take 7 to 9 months to hatch; the young begin to feed a month after hatching.The horn shark feeds on benthic invertebrates,including sea urchins (echinoids), crabs, shrimp, isopods, sipunculid worms,anemones, bivalves, gastropods (possibly abalone), cephalopods (octopuses), but less commonly on small fish includingpipefish (Syngnathidae) and blacksmith (Chromis punctipinnis, Pomacentridae). According to Michael (1993), the activediurnal blacksmith is eaten at night by the horn shark while it is resting on the bottom. Predators are little known: a Pacificangelshark (Squatina californica) has been filmed as swallowing small horn sharks and spitting them out alive, possiblybecause of their strong spines.Size: Maximum 122 cm but most adults are below 97 cm. Egg cases 10 to 12 cm long and 3 to 4 cm wide at broad end (notover flanges); length at hatching 15 to 16 cm; males maturing at about 58 to 59 cm and adult at 59 to 84 cm; females matureabove 58 cm.Interest to Fisheries and Human Impact: Interest to fisheries minimal, probably utilized or formerly utilized for fishmeal asa bycatch of the shrimp fishery and other bottom-trawling operations in Pacific Mexican waters. It has been captured bydivers for sport and for its large fin spines, which are made into jewellery; decreases in numbers of horn sharks have beennoted in areas with intense diver activity in southern California. Horn sharks are often harassed and grabbed by divers, butwhen provoked may swim after their assailants and bite them. These sharks are kept in many public aquaria in the UnitedStates. They are hardy, attractive, readily maintained, will breed in captivity, and have been displayed for many years.Local Names: California bull-head shark, Bullhead shark, Horned shark.Remarks: Michael (1993) had a photobraph and brief account of what may be an undescribed bullhead shark in thesouthern Gulf of California, which he termed the Cortez bullhead shark (Heterodontus sp.). According to Michael it is similarto H. francisci and H. mexicanus but differs from both species in having higher, more falcate dorsal fins, no dark spots, alighter abdomen, and no light line on the interorbital space. It has low supraorbital ridges as in H. mexicanus. The specieshas not, to the writer’s knowledge, been collected, so its status is treated as uncertain here pending detailed comparison ofmaterial with the sympatric H. mexicanus and H. francisci.Literature: Daniel (1928); Beebe and Tee-Van (1941); Smith (1942); Roedel and Ripley (1950); Limbaugh (1963); Nelsonand Johnson (1970); Miller and Lea (1972); Taylor (1972); Feder, Turner and Limbaugh (1974); Applegate et al. (1979);Chirichigno (1980); Compagno (1983, 1984); Michael (1993); Compagno, Krupp and Schneider (1995); Segura-Zarzosa,Abitia-Cárdenas and Galván-Magaña (1997).

38 FAO Species Catalogue for Fishery Purposes No. 1Heterodontus galeatus (Günther, 1870) Fig. 34Cestracion galeatus Günther, 1870, Cat. Fish. British Mus., 8: 416. Holotype: British Museum (Natural History), BMNH1862.7.2.2, about 64 cm long, catalogue number according to Eschmeyer (1998, Cat. Fish.: CD-ROM), Australia.Synonyms: None.Other Combinations: Gyropleurodus galeatus (Günther, 1870), Molochophrys galeatus (Günther, 1870).FAO Names: En - Crested bullhead shark; Fr - Requin dormeur à crête; Sp - Dormilón carenado.Field Marks: Dorsal fins with spines, anal fin present, supraorbital ridges greatly enlarged, colour pattern of dark broadbands on head, back and tail.Diagnostic Features: Supraorbital ridges very high, more prominent than in any other Heterodontus, abruptly truncatedposteriorly; interorbital space deeply concave, depth between ridges about equal to eye length. Anterior holding teeth with acusp and a pair of cusplets in adults, posterior molariform teeth strongly carinate and not greatly expanded and rounded.Pre-first dorsal-fin length 22 to 28%, and anal-caudal space 5 to 8%, of total length. Lateral trunk denticles fairly large andrough. Propterygium separate, not fused to mesopterygium. First dorsal-fin spine directed obliquely posterodorsally inadults; first dorsal-fin origin anterior to pectoral-fin insertions, slightly behind pectoral-fin midbases and well posterior to fifthgill openings; first dorsal-fin insertion well anterior to pelvic-fin origins and well behind pectoral-fin insertions; first dorsal-finfree rear tip about opposite to or slightly behind pelvic-fin origins; first dorsal fin moderately high and semifalcate or angularin adults, height 10.8 to 15.2% of total length, slightly larger than pelvic fins. Second dorsal-fin origin well in front of pelvic-finrear tips, second dorsal fin somewhat falcate or angular and nearly as large as first dorsal fin. Anal fin subangular or roundedto weakly falcate, apex reaching lower caudal-fin origin when laid back; anal-caudal space between 1 and 2 times anal-finbase. Total vertebral count 106 to 108, precaudal count 71 or 72, monospondylous precaudal count 34 or 35,diplospondylous precaudal count 36 to 38, pre-first dorsal-fin spine count 16 to 18, and count from diplospondyloustransition to second dorsal-fin spine 8 to 11. Egg cases with flat thin spiral flanges that are diagonal to case axis and a pair oflong, slender tendrils on case apex, flanges with 6 or 7turns. A large species, mature between 60 and 152 cm.Colour: background colour of dorsal surface light brownor yellowish brown with five broad diffuse-edged brown orblackish saddles, but without light or dark spots; saddlesnot arranged in a harness pattern; head with a dark baron interorbital surface and a single broad dark blotchunder eye; fins without abrupt dark tips and whitedorsal-fin apices; hatchlings without whorls on fins andbody, colour pattern as in adults.Distribution: Western South Pacific: East coast ofAustralia, from southern Queensland and New SouthWales, with a doubtful record from Tasmania, and apossible record from the top of the Cape York Peninsula.Fig. 34 Heterodontus galeatus

Sharks of the World, Vol. 2 39Habitat: A moderately common benthic and epibenthic shark of the southern Australian continental shelf at moderatedepths, ranging from close inshore in the intertidal zone to 93 m. Found on reefs, in kelp and in beds of seagrass.Biology: This shark often wedges its way between rocks in search of prey. The egg cases are dropped by females inseaweeds or sponges from 20 to 30 m depth on the bottom, during July and August, and hatch after about eight months.Eggs are commonest on the bottom in August and September but are found throughout the year. In captivity a newlyhatched female matured and began to lay eggs at an age of 11.8 years and a length of about 70 cm. The crested bullheadshark feeds primarily on sea urchins (echinoids), but also crustaceans, molluscs and small fishes.Size: Maximum said to be 152 cm long and attaining at least 130 cm, but most individuals are below 122 cm. Young hatch atabout 17 cm. Males mature at about 60 cm and females at about 70 cm.Interest to Fisheries and Human Impact: Interest to fisheries minimal, taken by bottom trawlers but not utilizedcommercially. Conservation status uncertain, distribution restricted to the warm east coast of Australia. Utilization byaquarium trade uncertain, but an obvious candidate because of its unusual appearance and striking colour pattern.Observed and photographed by divers, but not a special focus of ecotourism.Local Names: Crested Port Jackson shark, Crested shark, Crested horn shark.Literature: Whitley (1940); Fowler (1941); Smith (1942); McLaughlin and O’Gower (1971); Taylor (1972); Compagno(1984); Michael (1993); Last and Stevens (1994); Compagno and Niem (1998).Heterodontus japonicus (Maclay and Macleay, 1884) Fig. 35Cestracion japonicus Maclay and Macleay, 1884, Proc. Linnean Soc. New South Wales, 1884, 8(4): 428, pl. 20. Holotype:Australian Museum, Sydney, AMS B.68, female from Tokyo, Japan.Synonyms: ?Cestracion philippi var. japonicus Dumeril, 1865: 426. Dumeril proposed this as a colour variant of Squalusphilippi Bloch and Schneider, 1801 (= S. portusjacksoni Meyer, 1793) for Japanese specimens with transverse bands, butconsidered it equivalent to Centracion zebra Gray, 1831. Thus Dumeril may have confused two species, both of which occurin Japan. Taylor (1971: 107-108) requested the International Commission on Zoological Nomenclature to suppressDumeril’s name to remove the clearly defined Cestracion japonicus Maclay and Macleay, 1884, from possible juniorhomonymy if C. philippi var. japonicus is considered a synonym of C. zebra. Dumeril’s account could also be interpreted asproviding a separate name for a Japanese variety of C. philippi with transverse bands apart from C. zebra (which Dumerilalso synonymized with C. philippi), and recognized as defined by subsequent workers.Other Combinations: Gyropleurodus japonicus (Macleay and Macleay, 1884).FAO Names: En - Japanese bullhead shark; Fr - Requin dormeur nekozame; Sp - Dormilón japonés.Fig. 35 Heterodontus japonicus

40 FAO Species Catalogue for Fishery Purposes No. 1Field Marks: Dorsal fins with spines, anal fin present, first dorsal-fin origin over pectoral-fin bases, colour pattern of 11 to 14broad, irregular-edged, dark saddles and vertical stripes on a light background.Diagnostic Features: Supraorbital ridges moderately low, gradually ending posteriorly; interorbital space shallowlyconcave, depth between ridges about half eye length. Anterior holding teeth with a cusp and a pair of cusplets in adults,posterior molariform teeth not carinate and greatly expanded and rounded. Pre-first dorsal-fin length 21 to 25% andanal-caudal space 8 to 10% of total length. Lateral trunk denticles large and rough. Propterygium separate, not fused tomesopterygium. First dorsal-fin spine directed obliquely posterodorsally in hatchlings, juveniles and adults; first dorsal-finorigin anterior to pectoral-fin insertions and slightly behind pectoral-fin midbases, well posterior to fifth gill openings; firstdorsal-fin insertion well anterior to pelvic-fin origins, well behind pectoral-fin insertions; first dorsal-fin free rear tip aboutopposite to or slightly ahead or behind pelvic-fin origins; first dorsal fin very high and broadly semifalcate in young butmoderately high and semifalcate in adults, height 11 to 21% of total length, first dorsal fin much larger than pelvic fins.Second dorsal-fin origin over or slightly in front of pelvic-fin rear tips, broadly falcate and much smaller than first dorsal fin.Anal fin subangular and rounded to weakly falcate, apex well anterior to lower caudal-fin origin when laid back; anal-caudalspace nearly or quite twice anal-fin base. Total vertebral count 109 to 116, precaudal count 72 to 78, monospondylousprecaudal count 33 to 39, diplospondylous precaudal count 37 to 42, pre-first dorsal-fin spine count 15 to 17, and count fromdiplospondylous transition to second dorsal-fin spine 9 to 15. Egg cases with flat thin spiral flanges diagonal to case axis andhaving a pair of very short, slender tendrils on case apex, flanges with three turns. A large species, mature between 69 and120 cm. Colour: background colour of dorsal surface tan to brown with 11 to 14 brown diffuse-edged markings from snout tipto origin of caudal fin, including broad saddles and narrower vertical bands usually between them, body without light or darkspots, bands and saddles not arranged in a harness pattern; head with a light-coloured bar on interorbital surface, and with asingle broad dark blotch under eye that is indistinct in large adults; fins without abrupt dark tips and white dorsal-fin apices;hatchlings without whorls on fins and body, pattern as in adults although brighter.Distribution: Western North Pacific: Japan, Koreanpeninsula, northern China, and Taiwan (Province ofChina). An East African record is apparently erroneous.Habitat: A common, temperate-water bullhead shark ofthe western North Pacific continental shelf, occurring atmoderate depths of 6 to 37 m, on or near the bottom. Itprefers rocky areas (including reefs) and kelp-coveredbottom.Biology: This is a sluggish, slow-swimming shark, easilycaught by divers. It slowly explores the bottom, swimmingand ‘walking’ with its mobile paired fins.Oviparous, laying its large spiral-cased eggs among rocksor in kelp, at depths of about 8 or 9 m; several femalesmay lay their eggs in a single site, termed ‘nests’,although they apparently do not guard these sites after laying. In Japanese waters, eggs are laid from March throughSeptember, most abundantly in March through April; each female usually lays two eggs at a time, for 6 to 12 spawnings.Eggs hatch in about a year.The Japanese bullhead shark feeds on crustaceans, molluscs (including top shells [Trochidae; Gastropoda]), small fishesand sea urchins. It can protrude its jaws a considerable distance while grabbing prey.Size: Maximum total length about 120 cm. Size at hatching about 18 cm; males adult at 69 cm.Interest to Fisheries and Human Impact: Interest to fisheries probably minimal, caught and eaten in Japan andpresumably elsewhere in its range. Kept in public aquaria in Japan.Local Names: Bull head, Japanese bull-head shark, Cat shark, Japanese horn shark, Cestracion shark, Sazaewari,Sazaiwari, Nekozame (Japan); Japanse bulkophaai (South Africa).Literature: Fowler (1941); Smith (1942); Lindberg and Legeza (1959); Chen (1963); McLaughlin and O’Gower (1971);Taylor (1972); Nakaya and Shirai (1984); Compagno (1984); Michael (1993); Shen et al. (1993).

Sharks of the World, Vol. 2 41Heterodontus mexicanus Taylor and Castro-Aguirre, 1972 Fig. 36Heterodontus mexicanus Taylor and Castro-Aguirre, 1972, An. Esc. Nac. Cienc. Biol. México, 19: 125, figs 1-5, 8-9.Holotype: Scripps Institution of Oceanography, SIO-70-90, 610 mm adult female, Cerro Colorado, Sonora, Gulf of California,Mexico.Synonyms: None.Other Combinations: None.FAO Names: En - Mexican hornshark; Fr - Requin dormeur buffle; Sp - Dormilón búfalo.Fig. 36 Heterodontus mexicanusField Marks: Dorsal fins with spines, anal fin present, colour pattern of large dark spots of one-half eye diameter or more onlight background, a light bar present on space between supraorbital ridges, first dorsal-fin origin over pectoral-fin bases.Diagnostic Features: Supraorbital ridges low, gradually ending posteriorly; interorbital space shallowly concave, depthbetween ridges less than one-fourth eye length. Anterior holding teeth with a cusp and a pair of cusplets in adults, posteriormolariform teeth strongly carinate and not greatly expanded and rounded. Pre-first dorsal-fin length 24 to 29% andanal-caudal space 6 to 9% of total length. Lateral trunk denticles large and rough, area behind first dorsal fin with about 70 to130 denticles per cm 2 in adults. Propterygium fused to mesopterygium. First dorsal-fin spine directed obliquelyposterodorsally in juveniles and adults; first dorsal-fin origin slightly anterior to pectoral-fin insertions, behind pectoral-finmidbases, and well posterior to fifth gill openings; first dorsal-fin insertion well anterior to pelvic-fin origins and well behindpectoral-fin insertions; first dorsal-fin free rear tip about opposite to or slightly ahead of pelvic-fin origins; first dorsal fin lowand weakly falcate in adults, height 8 to 18% of total length, first dorsal fin about as large as pelvic fins; second dorsal-finorigin over or slightly in front of pelvic-fin rear tips, weakly falcate and nearly as large as first dorsal fin. Anal fin subangularand rounded to weakly falcate, with apex reaching lower caudal-fin origin or falling somewhat behind it when laid back;anal-caudal space between 1 and 2 times anal-fin base. Total vertebral count unknown, precaudal count 60 to 70,monospondylous precaudal count 30 to 34, diplospondylous precaudal count 30 to 38, pre-first dorsal-fin spine count 14 to16, and count from diplospondylous transition to second dorsal-fin spine 9 to 14. Egg cases with thick, T-shaped pairedspiral flanges, transverse to case axis, and a pair of long, slender tendrils on case apex; flanges with five turns. A smallspecies, mature between 50 and 70 cm. Colour: background colour of dorsal surface light grey-brown with large black spotson body and fins, these one-half eye diameter or more in size; body without a dark harness pattern; head with alight-coloured bar on interorbital surface of head and 1 or 2 dusky indistinct blotches under eye; fins without abrupt dark tipsand white dorsal-fin apices; hatchlings without whorls on fins and body.

42 FAO Species Catalogue for Fishery Purposes No. 1Distribution: Eastern Pacific: Mexico (southern BajaCalifornia, the Gulf of California, and southern Pacificcoast) south to Guatemala, Panama (Gulf of Panama),Colombia, probably Ecuador and Peru.Habitat: A warm-temperate and tropical bullhead sharkof littoral continental waters, found on rocky bottomincluding reefs and seamounts, on coral reefs, and onsandy areas from close inshore down to 20 to 50 mdepth.Biology: Common in the upper Gulf of California.Oviparous. The long tendrils and rigid, T-shaped spiralflanges on the egg cases of this shark suggest thatwedging of the eggs in crevices through the action offlexible flanges has been replaced by anchoring of thecases to the substrate by the tendrils, unlike otherbullhead sharks with flexible-flanged eggs. The heavyT-flanges may serve instead to protect the egg fromimpacts and egg-predators. Feeds on crabs anddemersal fishes including midshipman (Porichthys,Batrachoididae).Size: Maximum size about 70 cm. Egg cases about 8 to9 cm long, young hatch at about 14 cm; males maturingbetween 40 and 50 cm and reaching at least 55 cm.Interest to Fisheries and Human Impact: Interest to fisheries minimal. Small numbers are or were taken as a bycatch ofthe shrimp fishery in Mexico and processed into fishmeal along with other sharks. It is also caught in gill nets set for smallsharks. Observed by divers in the Gulf of California, but not a special focus for ecotouristic diving.Local Names: Buffalo hornshark, Mexican horn shark; Gata (Mexico).Remarks: This shark had been collected in the Gulf of California many years ago by ichthyologists at the Scripps Institutionof Oceanography and referred to under the unpublished manuscript names “H. seftoni” and “H. oligostictus”. It waseventually published as H. mexicanus.Literature: Taylor (1972); Taylor and Castro-Aguirre (1972); Applegate et al. (1979); Chirichigno (1980); Compagno (1984);Franke and Acero (1991); Michael (1993); Compagno, Krupp and Schneider (1995).Heterodontus portusjacksoni (Meyer, 1793) Fig. 37Squalus portus jacksoni Meyer, 1793, Syst. Summar. Zool. Entdeck. Neuholland, Afrika: 71. No type material, Botany Bay,New South Wales, Australia, based on the Port Jackson Shark of Phillip, 1789, Voyage Botany Bay: 283, fig.Synonyms: Squalus jacksoni Suckow, 1799: 102. No type material, Botany Bay, Port Jackson, Australia, based on the PortJackson Shark of Phillip, 1789: 283, fig. Reference from Fowler (1941). Squalus philippi Bloch and Schneider, 1801: 134.No type material, Botany Bay, Port Jackson, Australia, based on the Port Jackson Shark of Phillip, 1789: 283, fig. Squalusphilippinus Shaw, 1804: 341. No type material?, southern Pacific Ocean, Botany Bay (Port Jackson, Australia), apparentlybased on the Port Jackson Shark of Phillip (1789: 283, fig.), and termed the “Phillipian shark” by Shaw. Squalus jacksoniiTurton, 1806: 922. Variant spelling of Squalus jacksoni Suckow, 1799 or independently proposed? Cestracion philippiLesson, 1830, 2: 97; 3, pl. 2. No type material, Botany Bay, Port Jackson, Australia, based on the Port Jackson Shark ofPhillip, 1789: 283, fig. Proposed as a new name; specimen illustrated may not be this species. Cestracion heterodontusSherrard, 1896: 42, 88, figs. Hobson’s Bay, Victoria. Reference from Fowler (1941), uncertain if new name or error.Heterodontus bonae-spei Ogilby, 1908: 2. Holotype: Queensland Museum, No. QM I.1587, jaws only, “Table Bay, SouthAfrica”, possibly a specimen of H. portusjacksoni with a mistaken locality label according to Reif (1973: 165-167).Other Combinations: None.

Sharks of the World, Vol. 2 43FAO Names: En - Port Jackson shark; Fr - Requin dormeur taureau; Sp - Dormilón toro.Field Marks: Dorsal fins with spines, anal fin present, colour pattern with a conspicuous set of harness-like narrow darkstripes on the back, unique to the species.Diagnostic Features: Supraorbital ridges moderately low, gradually ending posteriorly; interorbital space moderatelyconcave, depth between ridges less than half eye length. Anterior holding teeth with a cusp and no cusplets in adults,posterior molariform teeth not carinate and greatly expanded and rounded. Pre-first dorsal-fin length 21 to 24% andanal-caudal space 10 to 13% of total length. Lateral trunk denticles fairly large and rough. Propterygium fused tomesopterygium. First dorsal-fin spine directed obliquely posterodorsally in juveniles and adults; first dorsal-fin origin wellanterior to pectoral-fin insertions, about over or slightly behind pectoral-fin midbases, and somewhat posterior to fifth gillopenings; first dorsal-fin insertion well anterior to pelvic-fin origins and well behind pectoral-fin insertions; first dorsal-fin freerear tip about opposite to pelvic-fin origins; first dorsal fin moderately high and rounded angular or falcate, height 12 to 16%of total length, first dorsal fin larger than pelvic fins; second dorsal-fin origin over or slightly behind pelvic-fin rear tips, seconddorsal fin rounded to angular or falcate and nearly as large as first dorsal fin. Anal fin subangular and rounded or weaklyfalcate, apex well anterior to lower caudal-fin origin when laid back; anal-caudal space about three times anal-fin base. Totalvertebral count 114, precaudal count 76 to 81, monospondylous precaudal count 37 to 39, diplospondylous precaudal count37 to 43, pre-first dorsal-fin spine count 15 to 17, and count from diplospondylous transition to second dorsal-fin spine 9 to14. Egg cases with flat thin spiral flanges diagonal to case axis and a pair of very short, slender tendrils on case apex;flanges with four or five turns. A large species, mature between 70 and 165 cm. Colour: background colour of dorsal surfacegrey to light brown or whitish with distinctive black striped harness marking; body and fins without light or dark spots; headwith a narrow dark bar on interorbital surface and a single narrow dark to blackish band under eye; fins without abrupt darktips and white dorsal-fin apices; hatchlings without whorls on fins and body, colour pattern as in adults.Distribution: Western South Pacific: Temperate andsubtropical southern Australia, from off New SouthWales, Victoria, Tasmania, South and Western Australia(west coast), with questionable records from southernQueensland and the tropical north coast of WesternAustralia; also New Zealand (a single record, possibly asa straggler or waif from Australia).Habitat: A common littoral, nocturnal bottom shark of thetemperate Australian continental shelves and uppermostslopes, ranging from close inshore in the intertidal to atleast 275 m.Fig. 37 Heterodontus portusjacksoniBiology: Underwater observation and tagging of thisnocturnal species has elucidated its life-history to adegree attained with few other species of sharks. Whileinshore, the Port Jackson shark favours caves withsandy floors and open trenches of shallow rocky reefs asdaytime resting places, and almost all individuals in a given area will be found resting in relatively few of such sites. Strongselection is shown for favoured sites, and superficially identical sites nearby may have few or no sharks.Port Jackson sharks can be solitary but often occur in small to large groups. Althose these sharks are evidently social andapparently are amenable to study underwater, relatively little is known of their sociobiology and behaviour patterns. Thiscould be elucidated in the future by electronic tagging and night-observation with low-light video devices, as well asobservations of captive colonies in semi-naturalistic habitats.

44 FAO Species Catalogue for Fishery Purposes No. 1Pronounced fluctuations in abundance have been noted on shallow reefs off New South Wales, directly correlated withseasonal influxes of adults for breeding and inversely correlated by seasonal variations in temperature. These sharks areapparently social while resting, and favoured resting sites may have up to 16 sharks occupying them. Data from taggingsuggests that seasonal reef populations are in a state of continuous flux, with individuals moving in and out of their favouredreefs throughout the breeding season. Apparently individuals are capable of homing to favoured resting sites after rangingconsiderable distances away from them during the breeding season. When sharks were experimentally removed fromresting sites in Sydney Harbour to different localities up to 3 km away, they returned to their original resting sites. It has beensuggested that these sharks have a highly-developed spatial memory, and apparently the means to locate favoured restingand breeding sites long distances apart along migration routes.Port Jackson sharks are seasonal oviparous breeders, with juveniles segregating by size after hatching and adultssegregating by sex. Mature females accompanied by some males move onto inshore reefs in late July and August in theSydney area (New South Wales), and probably mating occurs at this time. Most mature males remain in deeper wateroffshore. During August and September (rarely in July and October) females lay 10 to 16 (commonly 10 to 12) eggs in rockcrevices on shallow, sheltered reefs at depths from 1 to 5 m but occasionally down to 20 to 30 m. In captivity females lay apair of cased eggs a day every 8 to 17 days. The broad spiral flanges of the egg cases serve as anchors to keep themwedged in the rocks. Females apparently favour traditional ‘nest’ sites, which several apparently use collectively for manyyears. Apart from rock crevices, females may occasionally lay egg cases on open sand, and egg cases have been foundwedged under an underwater oil pipeline and in tin cans. Egg cases are oriented with their pointed ends into crevices, andfemales have been seen carrying egg cases, suggesting that females lay their eggs, pick them up at the broad end, andinsert them into appropriate crevices. According to Michael (1993), adults have been observed eating their own egg-cases(as in H. francisci).Young hatch after about 9 to 12 months and move into nursery areas in bays and estuaries. Some may retreat into deeperwater during summer, but most juveniles remain in mixed groups with a 1:1 sex ratio on the nursery grounds for severalyears. At the beginning of sexual maturity adolescents move into deeper water and segregate into male and female groups.After several years of adolescence, apparently spent at the outer edges of the continental shelves, these groups join theadult populations.Adult males apparently move into deeper water near the end of the breeding season, followed by the adult females in lateSeptember or October. Some adults move offshore into deeper water, but others migrate. Small numbers of adults mayreturn to the inshore breeding reefs as early as March or April of the next year, but most do not stay inshore and few sharksare present until the onset of the next breeding season. Observed ratios of adult males and females are not significantly atvariance with a 1:1 ratio.On the east coast of Australia the Port Jackson shark shows a pattern of migration southwards after breeding, with femalesmigrating at least for 5 to 6 months and moving up to 850 km south of breeding reefs before returning to the same sites thenext year. Some may range as far south as Tasmania from the Sydney area in New South Wales in the annual migrationcycle. It is thought that migrating adult sharks move southwards along inshore coastal waters but return to their breedingreefs along deeper offshore waters.Studies on blood proteins between Port Jackson sharks of different regions suggest that they form at least two populations,a southwestern one from Western Australia to northeastern Victoria and a northeastern one from New South Wales andpossibly southern Queensland. There is blood protein evidence to suggest that sharks using favoured breeding sites in threelocalities in New South Wales represent genetically distinct subpopulations, and indicates that the high site specificity shownby tagging and recapturing of sharks in this area is probably of relatively long duration.Data from captive sharks suggests that juveniles grow at about 5 to 6 cm per year and adults between 2 and 4 cm per year.Approximate estimates of age at maturity from captive growth data are 8 to 10 years for males and 11 to 14 years forfemales. So far, data is unavailable on growth rates in the wild from tagging and remeasuring of tagged individuals or fromcalibration and examination of fin spine or vertebral rings.The Port Jackson shark feeds primarily on benthic invertebrates, mainly echinoderms. Prey items include sea urchins,starfish, polychaetes, large gastropods, prawns, crabs, barnacles, and small fishes. Occasionally garbage such as bits ofmammalian fur, potato and orange peels are taken in by these sharks. Juveniles with their smaller, more pointed teethapparently take more soft-bodied prey than adults. Food items in stomachs are usually broken into small pieces, indicatingthat the sharks actively grind their food with their powerful jaws and heavy molariform teeth. Food is apparently taken at nighton the bottom, and by searching close to the substrate. Olfactory cues are thought to be important, but electrosense andlateral line sense may play a role in this also. Food is eaten after final contact with the mouth region. Juveniles at least arecapable of digging food out of the sand by sucking in water and sand and blowing it out of the gill covers. Respiration canoccur by pumping water into the first, enlarged gill slits and out the last four, which is thought to allow the shark to crush andgrind its prey at leisure without having to take in water through its mouth and risk passage of food out the gill slits.Predators of this shark are poorly known, but it is suspected that adults are highly protected by their sedentary habits,cryptic, nocturnal behaviour, fin spines, and disruptive colour patterns. Possible predators are large macropredatory sharkssuch as bluntnose sevengill and white sharks as well as large otariid seals. Juveniles in nursery grounds are thought to bemore vulnerable to predation by other sharks and larger benthic teleosts. Adults are sometimes attacked by small predatory

Sharks of the World, Vol. 2 45isopods, and eggs may be attacked by male Port Jackson sharks and possibly a gastropod drilling predator. As with othersharks, this has a sizeable parasite fauna, including cestodes (tapeworms), trematodes (flatworms), nematodes(roundworms), isopod larvae, copepods, fish lice, and leeches.Size: Maximum total length reported as 165 cm, but apparently rare above 137 cm. Egg cases are 13 to 17 cm long and 5 to7 cm wide at the broad end. Size at hatching 23 to 24 cm. Males are adolescent between 50 and 80 cm, mature between 70and 80 cm, and reach at least 105 cm; females are adolescent between 65 and about 84 cm, mature between 80 and 95 cm,and reach at least 123 cm; adult females average about 25 cm longer than adult males.Interest to Fisheries and Human Impact: Apparently of minimal interest to fisheries. Taken in commercial fisheries asbycatch in bottom trawls, shrimp nets, beach seines, anti-shark nets, bottom longlines and in shark gill nets on the southcoast of Australia; also caught by sports anglers on rod-and-reel. Apparently not utilized as food. This shark is consideredharmless to people. It is kept in public aquaria for display in Europe, the United States, and probably Australia, and is anobvious candidate for display because of its hardiness and attractive colour pattern. Divers observe this shark but it is not aspecial focus of ecotouristic diving. Conservation status uncertain.Local Names: Bullhead shark, Bullhead, Pigfish, Oyster-crusher or Oyster crusher, Tabbigaw.Remarks: Reif (1973) noted that the holotype of Heterodontus bonae-spei, supposedly from South Africa, is most probablya specimen of H. portusjacksoni with an erroneous locality label. Eschmeyer (1998) noted that the name was unavailablebecause Ogilby (1908) did not distinguish it by characters but only by locality.Literature: Ogilby (1908); Whitley (1940); Fowler (1941); Smith (1942); McLaughlin and O’Gower (1970, 1971); Taylor(1972); Reif (1973); O’Gower and Nash (1978); Michael (1993); Last and Stevens (1994); O’Gower (1995); Compagno andNiem (1998).Heterodontus quoyi (Fréminville, 1840) Fig. 38Cestracion quoyi Fréminville, 1840, Mag. Zool. Guerir., ser. 2(5): 1-3, pl. 3. Holotype: Museum National d’Histoire Naturelle,Paris, MNHN-3445, adult male about 475 mm, type locality Galapagos Islands.Synonyms: Cestracion pantherinus Valenciennes, 1846, pl. 10, fig. 2. Ibid., 1855, text: 350. Holotype the same specimen(MNHN-3445) as that of Cestracion quoyi, Galapagos Islands. Gyropleurodus peruanus Evermann and Radcliffe, 1917: 2,pl. 1, fig. 1. Holotype: U.S. National Museum of Natural History, USNM-77691, 565 mm TL adult (gravid) female, Lobos deTierra Island, Peru, confirmed by Howe and Springer (1993: 11).Other Combinations: None.FAO Names: En - Galapagos bullhead shark; Fr - Requin dormeur bouledogue; Sp - Dormilón de Galápagos.Fig. 38 Heterodontus quoyiField Marks: Dorsal fins with spines, anal fin present, first dorsal-fin origin over pectoral-fin inner margins, colour pattern oflarge dark spots.Diagnostic Features: Supraorbital ridges low, gradually ending posteriorly; interorbital space very shallowly concave,depth between ridges less than one-fourth eye length. Anterior holding teeth with a cusp and a pair of cusplets in adults,posterior molariform teeth strongly carinate and not greatly expanded and rounded. Pre-first dorsal-fin length 32 to 36% and

46 FAO Species Catalogue for Fishery Purposes No. 1anal-caudal space 4 to 7% of total length. Lateral trunk denticles fairly large and rough. Propterygium separate, not fused tomesopterygium. First dorsal-fin spine directed obliquely posterodorsally in hatchlings to adults; first dorsal-fin origin behindpectoral-fin insertions, over pectoral-fin inner margins and far behind gill openings; first dorsal-fin insertion about oppositepelvic-fin origins and far behind pectoral-fin insertions; first dorsal-fin free rear tip over or behind midbases of pelvic fins andsometimes about opposite pelvic-fin insertions; first dorsal fin rounded and brush-shaped in young and low androunded-subangular in adults, height 8 to 9% of total length, first dorsal fin subequal to pelvic fins; second dorsal-fin originslightly to well behind pelvic-fin free rear tips, second dorsal fin rounded-angular and nearly as large as first dorsal fin. Analfin rounded-angular, apex well anterior or reaching lower caudal-fin origin when laid back; anal-caudal space less than twiceanal-fin base. Total vertebral count 103 to 109, precaudal count 67 to 72, monospondylous precaudal count 24 to 36,diplospondylous precaudal count 33 to 41, pre-first dorsal-fin spine count 19 to 20, and count from diplospondyloustransition to second dorsal-fin spine 11 to 19. Identification of egg cases uncertain, but possibly like those of H. francisci,with flat thin spiral flanges diagonal to case axis, without tendrils on case apices, and flanges with five turns. A small species,mature between 48 and 61 cm. Colour: background colour of dorsal surface light grey or brown with large black spotsgreater than half eye diameter, no dark harness pattern; head without a light-coloured bar on interorbital surface and withmottled dark spots or blotches under eye; fins without abrupt dark tips and white dorsal-fin apices; hatchlings without whorlson fins and body and similar in coloration to adults.Distribution: Eastern Pacific from the coasts andoffshore islands of Peru and the Galapagos Islands.Habitat: A little-known but apparently common tropicaland warm-temperate bullhead shark of inshorecontinental and insular waters, at moderate depths onthe bottom. Lives on rocky and coral reefs, often seenresting on ledges of vertical rock surfaces at 16 to 30 mdepth.Biology: A poorly known, primarily nocturnal shark.Oviparous. Feeds on crabs; sometimes with marinealgae in its stomach. One taken from the stomach of atiger shark.Size: Maximum total length 61 cm; an egg case possiblyfrom this species was about 11 cm long; an apparentlynewly hatched male was 17 cm and an adult male was48 cm long.Interest to Fisheries and Human Impact: Not a commercial species (N. Chirichigno, pers. comm.), though presumablycaught as discarded bycatch. Commonly seen by divers off the Galapagos Islands.Local Names: Galapagos bull-head shark, Peruvian horn shark, Galapagos horn shark (English); Gato, Suño, Tiburóntamborín (Peru).Remarks: N. Chirichigno (1980, pers. comm. to Compagno, 1984) suggested that there may be more than one speciesincluded under H. quoyi. The quoyi-like Heterodontus from Peru, with the first dorsal-fin origin slightly behind thepectoral-fin bases, includes two forms: one of these has concave posterior dorsal-fin margins, a long space about twice theanal-fin base length between the anal-fin base and lower caudal-fin origin, and an anal fin that falls well ahead of the lowercaudal-fin origin when laid back; and a second form with convex posterior dorsal-fin margins, a short space much less thantwice the anal-fin base length between the anal-fin base and lower caudal-fin origin, and an anal fin that reaches the lowercaudal-fin origin when laid back. If distinct species, the first type is apparently the true H. quoyi, while the second could bedistinguished as H. peruanus. I continue to hesitate to separate these two forms with the small amount of material I haveexamined, and follow Taylor (1972), who examined material from Peru and included them in one species.Literature: Beebe and Tee-Van (1941); Smith (1942); McLaughlin and O’Gower (1971); Taylor (1972); Chirichigno (1980);Compagno (1984); Michael (1993); Compagno, Krupp, and Schneider (1995).Heterodontus ramalheira (Smith, 1949) Fig. 39Gyropleurodus ramalheira Smith, 1949a, Ann. Mag. Nat. Hist. (ser. 12), 2(17): 367, fig. 1. Holotype in Natural HistoryMuseum, Maputo, Mozambique, 585 mm female, moderately deep water off Inhambane, Mozambique.Synonyms: None.Other Combinations: None.

Sharks of the World, Vol. 2 47FAO Names: En - Whitespotted bullhead shark; Fr - Requin dormeur chabot; Sp - Dormilón boquigrande.Field Marks: Dorsal fins with spines, anal fin present, colour pattern of white spots on variegated darker backgroundincluding dark saddles in adults, hatchling young with whorls of dark lines on a light background.Diagnostic Features: Supraorbital ridges moderately high, abruptly truncated posteriorly; interorbital space moderatelyconcave, depth between ridges about half eye length. Anterior holding teeth with a cusp and a pair of cusplets in adults,posterior molariform teeth strongly carinate and not greatly expanded and rounded. Pre-first dorsal-fin length 20 to 26% andanal-caudal space 8 to 10% of total length. Lateral trunk denticles large and rough. Propterygium separate, not fused tomesopterygium. First dorsal-fin spine directed somewhat forward in hatchlings and juveniles and vertical in adults; firstdorsal-fin origin far anterior to pectoral-fin insertions, just behind or even over pectoral-fin origins and over third to fifth gillopenings; first dorsal-fin insertion far anterior to pelvic-fin origins, just behind pectoral-fin insertions; first dorsal-fin free reartip anterior to or opposite of pelvic-fin origins; first dorsal fin falcate in young and high and semifalcate in adults, firstdorsal-fin height 11 to 21% of total length, first dorsal fin much larger than pelvic fins; second dorsal-fin origin over pelvic-fininner margins and well in front of pelvic-fin rear tips, second dorsal fin falcate and much smaller than first dorsal fin. Anal finangular and falcate, apex slightly anterior to lower caudal-fin origin when laid back; anal-caudal space slightly less thantwice anal-fin base. Total vertebral count 104 to 116, precaudal count 67 to 73, monospondylous precaudal count 32 to 34,diplospondylous precaudal count 34 to 40, pre-first dorsal-fin spine count 10 to 14, and count from diplospondyloustransition to second dorsal-fin spine 7 to 10. Egg casesunknown. A moderately large species, mature between60 and 83 cm. Colour: background colour of dorsalsurface dark reddish brown with white spots, lighter inhatchlings, without a dark harness pattern but withdarker indistinct saddles; head without light-coloured baron interorbital surface of head in adults but young withtransverse parallel dark lines there, and a series ofnarrow dark parallel stripes under eye in hatchlings,changing to a dusky patch in larger juveniles and lost inadults; fins without abrupt dark tips and white dorsal-finapices; hatchlings with a unique and striking pattern ofnumerous thin curved parallel dark lines in whorls on finsand body, lost with growth and absent in adults.Distribution: Western and northern Indian Ocean,South Africa (KwaZulu-Natal), south-centralMozambique, Somalia, eastern shore of the ArabianPeninsula and southern Oman.Habitat: A rare and little-known benthic shark of theouter continental shelf and uppermost slope of southernand East Africa and the eastern Arabian Peninsula,unusual for the family in being a deepish water speciesfound at 40 to 275 m, with most records below 100 m andfrom trawler hauls. At least one station that recorded thisshark was on sandy bottom.Fig. 39 Heterodontus ramalheira

48 FAO Species Catalogue for Fishery Purposes No. 1Biology: Presumably oviparous, but egg cases have not been reported to date. Young individuals including a hatchlinghave been found off southern Mozambique at 110 m. Crabs were found in the stomachs of two individuals.Size: Maximum about 83 cm; hatchling 18 cm; males immature at 39 cm, adolescent at 56 cm, adults to at least 69 cm; adultfemales 75 to 83 cm.Interest to Fisheries and Human Impact: Interest to fisheries none, occasionally caught as bycatch of commercial bottomtrawlers including shrimp trawlers off southern Mozambique and South Africa. Conservation status unknown, apparentlyrare or uncommon, only one specimen caught recently in experimental trawling off Mozambique (Sea Fisheries ResearchInstitute, R.V. ALGOA cruise 014, 1994) with 52 offshore bottom trawl stations at depths of 37 to 517 m.Local Names: Mozambique bullhead shark, Mosambiekse bulkophaai (South Africa); Turbarão dorminhoco deMoçambique.Literature: Smith (1949a); Pinchuk (1969); Taylor (1972); Bass, D’Aubrey and Kistnasamy (1975d); Compagno (1984); vander Elst and Vermeulen (1986); Compagno, Ebert and Smale (1989); Bass (1986); Randall (1995); S. Dudley and P. vanBlerck (pers. comm.).Heterodontus zebra (Gray, 1831) Fig. 40Centracion zebra Gray, 1831, Zool. Misc.: 5. Holotype: British Museum (Natural History), BMNH 1953.5.10.4, dryspecimen, female about 47 cm, from Swatow, China (confirmed by Eschmeyer, 1998, Cat. Fish., CD-ROM, who gives thecatalogue number). Also, Cestracion zebra Agassiz, 1853, Proc. Am. Acad. Sci., 3: 65 (Eschmeyer, ibid.), possibly acorrection of Gray’s generic allocation rather than a new name.Synonyms: ?Cestracion philippi var. japonicus Dumeril, 1865: 426 (In part? See note above under H. japonicus).Cestracion amboinensis Regan, 1906b: 436. Holotype: British Museum (Natural History), BMNH 1867.11.28.100 or 183,580 mm specimen, Amboyna (confirmed by Eschmeyer, 1998: CD-ROM, who gives the catalogue number).FAO Names: En - Zebra bullhead shark; Fr - Requin dormeur zebre; Sp - Dormilón acebrado.Fig. 40 Heterodontus zebraField Marks: Dorsal fins with spines, anal fin present, striking zebra-striped colour pattern of numerous narrow dark verticalsaddles and bands on light background.Diagnostic Features: Supraorbital ridges low, gradually ending posteriorly; interorbital space very shallowly concave withdepth between ridges about one-fourth eye length. Anterior holding teeth with a cusp and probably a pair of cusplets inadults, posterior molariform teeth strongly carinate and not greatly expanded and rounded. Pre-first dorsal-fin length 21 to

Sharks of the World, Vol. 2 4927% and anal-caudal space 10 to 12% of total length. Lateral trunk denticles fairly small and smooth. Propterygium fused tomesopterygium. First dorsal-fin spine directed obliquely posterodorsally in young and adults; first dorsal-fin origin anterior topectoral-fin insertions, slightly behind pectoral-fin midbases, and well posterior to fifth gill openings; first dorsal-fin insertionwell anterior to pelvic-fin origins and well behind pectoral-fin insertions; first dorsal-fin free rear tip about opposite to orsomewhat behind pelvic-fin origins; first dorsal fin very high and falcate in young and moderately high and falcate in adults,first dorsal-fin height 9 to 27% of total length, first dorsal fin much larger than pelvic fins; second dorsal-fin origin behindpelvic-fin rear tips, second dorsal fin falcate and much smaller than first dorsal fin. Anal fin subangular and rounded tofalcate, apex slightly anterior to lower caudal-fin origin when laid back; anal-caudal space over twice anal-fin base. Totalvertebral count 117, precaudal count 74 to 81, monospondylous precaudal count 34 to 38, diplospondylous precaudal count39 to 45, pre-first dorsal-fin spine count 15 to 17, and count from diplospondylous transition to second dorsal-fin spine 10 to16. Egg cases with flat thin spiral flanges nearly transverse to case axis, without tendrils on case apices but with short oneson opposite end, flanges with a single turn. A large species, mature between 64 and 122 cm. Colour: background colour ofdorsal surface white or cream with a zebra-striped pattern of 22 to 36 brown or black, narrow vertical markings from snout tipto origin of caudal fin, with bold saddles and bands often separated by more diffused narrow bands, without light or darkspots, bands not arranged in a harness pattern; head with transverse dark and light bars on interorbital surface, and with abilobate pair of dark bands separated by a light stripe under eye; fins without abrupt dark tips and white dorsal-fin apices;hatchlings without whorls on fins and body, colour pattern as in adults.Distribution: Western Pacific: Japan, Korean peninsula,China, Taiwan (Province of China), Viet Nam, Indonesia(Sulawesi, Ambon), and tropical Australia (northernWestern Australia).Habitat: A common but little-known bottom shark, foundon the continental and insular shelves of the westernPacific from inshore down to at least 50 m in the SouthChina Sea, but deeper and in 150 to 200 m off WesternAustralia.Biology: Oviparous, details of spawning not recorded.Biology poorly known. Probably eats bottominvertebrates as with other members of the family.Size: Maximum total length about 122 cm, hatchlings atleast 15 cm, males immature at 44 cm and mature at 64to 84 cm, females to 122 cm.Interest to Fisheries and Human Impact: Probably ofminimal interest to commercial fisheries. Caught asbycatch by commercial trawlers and possibly otherfisheries in its range. Conservation status unknown but ofsome concern. Utilization in aquarium trade notrecorded, but an obvious candidate because of itsattractive colour pattern.Local Names: Zebra horn shark, Zebra Port Jacksonshark, Striped bullhead shark, Barred bull-head shark,Barred shark, Striped cat shark, Shima-nekozame(Japan); Maou urh sha or Cat shark, Mau i sha or Littleshark (China).Literature: Fowler (1941); Smith (1942); Lindberg and Legeza (1959); Chen (1963); Bessednov (1969); Taylor (1972);Compagno (1984); Nakaya and Shirai (1984); Last and Stevens (1994); Compagno and Niem (1998).Heterodontus sp. A Fig. 41Heterodontus sp. A J. Mee, pers. comm. for a bullhead shark collected off Oman, apparently representing an undescribedspecies. Also Randall, 1995, Coastal Fishes of Oman: 19, fig. 2.Synonyms: None.Other Combinations: None.

50 FAO Species Catalogue for Fishery Purposes No. 1FAO Names: En - Oman bullhead shark; Fr - Requin dormeur d’Oman; Sp - Dormilón de Omán.Field Marks: Dorsal fins with spines, anal fin present, first dorsal-fin origin over pectoral-fin inner margins, colour pattern of afew broad dark saddles on body, without narrow intermediate stripes or small dark spots, dorsal, caudal and pectoral finswith abruptly black tips, dorsal fins with white apical spots.Diagnostic Features: Supraorbital ridges moderately low, gradually ending posteriorly. Interorbital space weakly concave,depth between ridges probably less than one-fourth eye length. Anterior holding teeth with a cusp and a pair of cusplets inadults, posterior molariform teeth unknown. Pre-first dorsal-fin length about 24% and anal-caudal space about 7% of totallength. Lateral trunk denticles fairly rough. Propterygium condition unknown. First dorsal-fin spine directed obliquelyposterodorsally in adults, first dorsal-fin origin in front of pectoral-fin insertions, somewhat behind pectoral-fin midbases, andwell posterior to fifth gill openings; first dorsal-fin insertion well anterior to pelvic-fin origins and well behind pectoral-fininsertions; first dorsal-fin free rear tip slightly anterior to pelvic-fin origins; first dorsal fin low and rounded-angular in adults,height about 11% of total length, first dorsal fin about as large as pelvic fins; second dorsal-fin origin about opposite pelvic-fininsertions or rear tips, second dorsal fin weakly falcate and nearly as large as first dorsal fin. Anal fin subangular, with apexreaching lower caudal-fin origin when laid back; anal-caudal space less than twice anal-fin base. Total vertebral count 106,precaudal count 68, monospondylous precaudal count 35, diplospondylous precaudal count 33, prefirst dorsal-fin spinecount 13, and count from diplospondylous transition to second dorsal-fin spine 8. Egg cases unknown. Possibly a smallspecies, mature between 52 and 61 cm. Colour: background colour of dorsal surface tan to brown with 4 or 5 broaddiffuse-edged brown saddles from snout tip to origin of caudal fin, without light or dark spots except dark fin tips, no harnesspattern; head with a dark bar on interorbital surface and a single broad dark blotch under eye; fins tipped with dark brown orblackish, additionally a white spot on apices of dorsal fins; hatchling colour pattern unknown.Distribution: Northern Indian Ocean: Oman.Habitat: Caught off southern Oman, depth 80 m,presumably on soft bottom as specimens came fromcommercial trawlers.Biology: Essentially unknown.Size: Maximum size 61 cm. An adult male is 52 cm andan adult female 61.2 cm.Interest to Fisheries and Human Impact: Caught inhauls by commercial trawlers off southern Oman.Local Names: Oman bullhead shark.Remarks: This is apparently an undescribed species ofbullhead shark, superficially similar to H. japonicus butreadily separable by its different colour pattern, very lowdorsal fins, and possibly smaller size.Literature: Randall (1995); J. Mee (pers. comm).Fig. 41 Heterodontus sp. Aclick for next page

click for previous pageSharks of the World, Vol. 2 512.2 Order LAMNIFORMES - Mackerel sharksOrder: Group Lamnae Garman, 1885, Bull. Mus. Comp. Zool. Harvard, 12(1): 30. Emended here to Order LamniformesGarman, 1885.Number of Recognized Families: 5.Synonyms: Part 1 Squali, Abtheilung 2: Müller and Henle, 1838d: 27; Müller and Henle, 1839: 27. Part 1 Squali, Abtheilung 2,Unterabtheilung 3: Müller and Henle, 1839: 66. Ordo Plagiostomi, Subordo Squalini, Sectio Proktopterides, Tribus Dinotopterini:Bleeker, 1859: xi. Order Squali, Suborder Squali: Gill, 1862b: 394, 396. Order Squali, Suborder Galei: Gill, 1872: 22, 23. OrderPlagiostomi diplospondyli, Suborder Plagiostomi asterospondyli, Group 2 Scylliolamnidae: Hasse, 1879: 51. Order Selachii,Suborder Asterospondyli: Woodward, 1889: 157. Order Asterospondyli: Gill, 1893: 130; Fowler, 1941: 4, 13; Smith, 1949: 37, 39.Order Asterospondyli, Suborder Galei: Jordan and Evermann, 1896: 19, 21. Order Euselachii, Suborder Pleurotremata, DivisionGaleoidei: Regan, 1906: 723. Order Selachii, Group 2, Division B, Subdivision 1, Suborder Scylliodei: Goodrich, 1909: 148. OrderPleurotremata, Suborder Galeoidei: Engelhardt, 1913: 97. Order Plagiostoma, Suborder Antacea, Group Carcharoidei: Garman,1913: 10, 11. Order Plagiostoma, Suborder Antacea, Group Isuroidei: Garman, 1913: 10, 12. Order Euselachii, Suborder Galei,Series Lamnoidei: Jordan, 1923: 99. Order Plagiostomi, Suborder Galeiformes: Lozano y Rey, 1928: 280. Order Galea, SuborderIsurida, Superfamily Odontaspoidea: White, 1936: 4; White, 1937: 36, tab. 1. Order Galea, Suborder Isurida, Superfamily Isuroidea:White, 1936: 4; White, 1937: 36, tab. 1. Order Euselachii, Suborder Lamniformes: Bertin, 1939a: 9. Order Lamniformes: Berg, 1940:137; Berg and Svetovidov, 1955: 65; Patterson, 1967: 670; Rass and Lindberg, 1971: 303; Lindberg, 1971: 8, 257; Compagno,1973: 28; Applegate, 1974: 743; Nelson, 1976: 33; Compagno, 1984: 212; Nelson, 1984: 51; Gubanov, Kondyurin and Myagkov,1986: 3, 49; Cappetta, 1987: 26, 85; Compagno, 1988: 382; Eschmeyer, 1990: 435; Nelson, 1994: 51; de Carvalho, 1996: 55;Shirai, 1996: 32; Eschmeyer, 1998. Order Lamniformes, Suborder Lamnoidei: Berg, 1940: 137; Berg and Svedovidov, 1955: 65;Patterson, 1967: 670; Lindberg, 1971: 8, 257; Nelson, 1976: 33; Nelson, 1984: 51. Order Euselachii, Suborder Galei, SuperfamilyOdontaspoidea: Whitley, 1940: 68. Order Euselachii, Suborder Galei, Superfamily Isuroidea: Whitley, 1940: 68. Order Selachii,Suborder Galeoidea: Romer, 1945: 576; Bigelow and Schroeder, 1948: 77, 95; Romer, 1966: 350. Order Lamnoidea, SuborderGaleoidea: Schultz and Stern, 1948: 224. Order Lamnida, Suborder Lamnina: Matsubara, 1955: 1-789. Order Galeiformes,Suborder Isuroidea: Arambourg and Bertin, 1958: 2029. Order Pleurotrema, Suborder Galeoidea: Norman, 1966: 7. OrderCarchariida, Suborder Carchariina, Superfamily Carchariicae: Fowler, 1967a: 92, 140. Order Carchariida, Suborder Carchariina,Superfamily Lamnicae: Fowler, 1967a: 92, 104. Order Squatinida, Suborder Squaloidei: Glikman, 1967: 215. Superorder Lamnae,Order Odontaspidida: Glikman, 1967: 229, 230. Order Odontaspidida, Superfamily Odontaspidoidea: Glikman, 1967: 230. OrderOdontaspidida, Superfamily Isuroidea: Glikman, 1967: 232. Order Odontaspidida, Superfamily Scapanorhynchoidea: Glikman,1967: 233. Order Euselachii, Suborder Galeoidei: Blot, 1969: 702-776. Order Pleurotremata, Suborder Galeiformes: Budker andWhitehead, 1971: 5, tab. 2. Order Carcharhiniformes: Rass and Lindberg, 1971: 303; Gubanov, Kondyurin and Myagkov, 1986: 3,61. Order Isuriformes: Chu and Meng, 1979: 114, tab. 2. Order Isuriformes, Suborder Carcharioidea: Chu and Meng, 1979: 114, tab.2. Order Isuriformes, Suborder Isuroidea: Chu and Meng, 1979: 114, tab. 2. Order Isuriformes, Suborder Cetorhinoidea: Chu andMeng, 1979: 114, tab. 2. Order Isuriformes, Suborder Alopioidea: Chu and Meng, 1979: 114, tab. 2. Order Galeomorpha, SuborderLamnoidea: Carroll, 1988: 599.FAO Names: En - Mackerel sharks.Field Marks: Large active pelagic sharks without nictitating eyelids, no barbels or nasoral grooves, nostrils free from mouth,long mouths that extend behind eyes, usually with enlarged anterior teeth and a gap or small intermediate teeth betweenanteriors and laterals on each side of the upper jaw, five broad gill openings, two spineless dorsal fins and an anal fin.Diagnostic Features: Head conical to moderately depressed, not expanded laterally. Snout very short to moderatelyelongated, truncated to conical or blade-like and flattened, not greatly elongated and without lateral teeth or rostral barbels.Eyes usually on sides of head (dorsolateral in Carcharias), without nictitating lower eyelids, secondary lower eyelids, orsubocular pouches; upper eyelids not fused to eyeball. Nostrils of the ordinary shark type, transverse on snout, withoutbarbels, nasoral grooves or circumnarial grooves, separate from mouth, anterior nasal flaps short and not reaching mouth.Five pairs of gill openings present on sides of head, with the posteriormost two in front of pectoral-fin origins or above them.Spiracles present and very small, well behind and about opposite to level of eyes. Mouth large, arched and elongated,extending well behind eyes. Labial furrows reduced or absent, when present on both jaws or on the lower jaw only. Teethweakly to strongly differentiated along the jaws, with or without (Megachasma) enlarged anterior teeth but without enlargedmolariform posterior teeth; usually with a gap or small intermediate teeth between anterior and lateral teeth in the upper jaw(absent in Megachasma); teeth with osteodont histological structure. Trunk cylindrical, fusiform, or somewhat compressed,not flattened and ray-like. Caudal peduncle without thin lateral dermal ridges but with lateral keels variably present orabsent. Dermal denticles covering entire body, not enlarged as thorns or spines. Pectoral fins small to moderately large, notexpanded and ray-like, without triangular anterior lobes that cover the gill slits. Pectoral girdle (scapulocoracoid) high,U-shaped, with or (usually) without a medial joint, and with superscapulae directed posterodorsally and not contactingvertebral column. Pectoral-fin skeleton primitively tribasal, with propterygium in contact with radials and metapterygiumwithout a proximal segment; pectoral fins primitively aplesodic, with radials confined to the fin bases, but plesodic in derivedtaxa and supporting the fin webs; radial count 15 to 46 with 2 to 13 segments. Pelvic fins small to moderately large, with ventcontinuous with their inner margins. Claspers with siphons in the abdomen but without large clasper sacs; clasper glans witha pseudosiphon, cover rhipidion, rhipidion (sometimes absent), and clasper spurs or spines; dorsal and ventral marginals ofclasper skeleton rolled into a tube for the clasper canal. Two spineless dorsal fins present, with origin of first over abdomenand well in front of pelvic-fin origins; dorsal-fin skeleton with segmented radials but without segmented basal plates. Anal fin

52 FAO Species Catalogue for Fishery Purposes No. 1present. Caudal fin with a long dorsal lobe and the ventral lobe very strong to absent; vertebral axis elevated into the dorsalcaudal lobe (heterocercal caudal fin). Vertebral calcification usually strong, secondary calcification usually in form of strongbranched radii in intermedial spaces, sometimes with annular rings but without diagonal lamellae or other calcifications inthe basal spaces (Megachasma with vertebral calcification greatly reduced). Total vertebral count 109 to 477, precaudalvertebrae 50 to 125. Neurocranium with a short to greatly elongated, tripodal rostrum without ventral keel and open dorsally;nasal capsules spherical, oval or flattened and without subnasal fenestrae (basal communicating canals) or antorbitalcartilages; orbits with complete preorbital walls (except Mitsukurina), strong supraorbital crests (reduced to isolatedpreorbital and postorbital processes in Mitsukurina), strong suborbital shelves, separate foramina for superficialophthalmic nerves and hyomandibular nerves, and incomplete postorbital walls without lateral commissures for lateral headvein; occipital condyles low, occipital hemicentrum present between them. Jaws elongated, upper jaws (palatoquadrates)with or without short vertical orbital processes that articulate with cranial orbits in orbital notches of suborbital shelves or withbasal plate; orbital processes when present do not penetrate supraorbital crests. Hyobranchial skeleton with narrowelongated basihyoid; posterior two pharyngobranchials and last epibranchial fused into a yoke-shaped element. Headmuscles include elongated horizontal preorbitalis, elongated broad levator palatoquadrati that extend far behind the orbits,adductor mandibulae muscles not segmented and notched anteriorly for mouth gape; no craniomandibular muscle betweenthe lower jaw and orbital walls; no mandibulocutaneous muscle between upper jaw and skin; and no postocular eyelidmuscles. Intestinal valve of ring type, with 19 to 55 turns. Reproduction ovoviviparous (aplacental viviparous), foetalnutriment in at least some species from uterine cannibalism (eating of eggs and, in Carcharias taurus at least, eating otherfoetuses), but without placental vivipary or nutritive trophonemata.Distribution: Circumglobal in temperate and tropical seas, with some lamnoids penetrating cold boreal and subantarcticwaters. Some species favour temperate to cold boreal or subantarctic waters (basking shark, porbeagle and salmon shark)while most other species occur in warm-temperate to tropical seas; the white shark has one of the most extensive ranges ofany cartilaginous fish.Habitat: Mackerel sharks or lamnoids occur in a variety of marine habitats from shallow open and enclosed bays, rocky andcoral reefs, and sandy beaches on the continental shelves to the epipelagic zone and possibly the mesopelagic zone of theopen ocean, with a few species occurring on the continental and insular slopes. They range in depth from the intertidal to atleast 1 600 m on the lower slopes, in the open ocean from the surface to at least 450 m, and on the abyssal plains overbottoms down to over 5 000 m depth. Although a few species are found in shallow bays and off beaches in the intertidal zonelamnoid sharks are not known to penetrate brackish estuaries and are not recorded from freshwater rivers and lakes.Biology: The lamnoids are a small group of possibly 15 living species but are remarkably varied and often specialized intheir form and habits. Except for the smallish crocodile shark, all living lamnoids are medium-sized to gigantic. Their ranksinclude sharks of littoral morphotype (Compagno, 1990a: sand tiger shark, Carcharias taurus), but also high-speedtachypelagic predators (shortfin mako), macroceanic and microceanic specialists of the open ocean (longfin mako andcrocodile shark), two very different filter feeders (basking and megamouth sharks), bathic and rhynchobathic deepwaterspecialists (bigeye sand tiger and goblin sharks), and an archipelagic shark or top predator (white shark). There are no livingdurophage (shell-crushing) or specialized bottom-dwelling lamnoids. The prey range of lamnoids is vast, from microscopiczooplankton to large bony fishes, marine mammals, other chondrichthyans, marine birds and reptiles, cephalopods, crabs,large gastropods, and carrion. Most lamnoids are active swimmers, some are highly migratory, and some may seasonallyvisit favoured areas including concentrations of food (fish banks, areas and current systems with plankton blooms, and sealcolonies). At least some of the species are social (sand tiger shark, Lamnidae), and some may practice cooperative hunting.Mode of reproduction is known for only some of the lamnoid species which practice uterine cannibalism in the form ofegg-eating or oophagy (Lamnidae, Alopiidae, crocodile shark, sand tiger shark, and possibly basking and megamouthsharks), but also foetus-eating or adelphophagy (sand tiger shark, possibly crocodile shark).Interest to Fisheries and Human Impact: Several are important fisheries sharks in coastal and oceanic waters,particularly members of the families Alopiidae, Cetorhinidae, Lamnidae, and some Odontaspididae. These are regularcomponents of targeted commercial or sport fisheries including some fisheries specifically targeting certain species(basking, shortfin mako and white sharks), and as bycatch of other fisheries targeting teleost fishes or marine invertebrates.Some species are rare to common and often discarded bycatch of high-seas and deepwater fisheries (crocodile shark,bigeye sand tiger, goblin shark). Lamnoids are caught in bottom and pelagic trawls, in pelagic and fixed gill nets (includinganti-shark nets to protect bathing beaches), in fish traps, on bottom and pelagic longlines, in purse seines, with harpoons,and with hook-and-line and rod-and-reel. Many species are used for human consumption; the flesh of some species isexcellent, and large fins are of high value in the oriental soup-fin trade. Several inshore and offshore species are caught bysportsfishing anglers, and some species (makos, white shark, porbeagle, threshers) are sought by big-game anglers andare recognized by the International Game Fish Association.White sharks rarely but regularly bite swimmers, surfers, divers and boats, but less than a third of such incidents result infatalities and very rarely result in consumption of the victim. Sand tigers and shortfin makos sporadically bite but do notconsume people, and shortfin makos sometimes cause problems by jumping into sportsfishing boats after being hooked byanglers. Jaws and more recent shark-monster Hollywood films have inspired social phenomena leading to conservation problemsfor white sharks and sharks in general but have also been instrumental in promoting public awareness of the issues of sharkconservation. Hollywood lamnoid stars have suffered from increased fishing pressure and demand for trophies and food includingjaws, teeth, meat and fins, but have also received public support for their protection and conservation.

Sharks of the World, Vol. 2 53White sharks, shortfin makos, sand tiger sharks, and even the smalltooth sandtiger (Odontaspis ferox) are currently soughtby ecotouristic divers and film-makers in the Indian Ocean, western Atlantic, western Pacific and eastern Pacific. Althoughthe white shark has been subject to repeated and unsuccessful attempts to keep it in captivity in large public aquaria, thesand tiger shark is the only lamnoid that is readily kept for public viewing and is currently living in numerous aquaria in theUSA, Europe and South Africa.There are conservation concerns over several lamnoid species, including the white, basking and sand tiger sharks, some ofwhich have declined locally from overfishing and possibly other environmental problems. Certain species yield products ofextremely high value comparable to rhinoceros horns and elephant tusks, including the jaws, teeth and fins of white sharksand the fins of basking sharks. More abundant oceanic fisheries species, including makos and threshers, are in need ofrational management, while less abundant to rare offshore species such as crocodile and bigeye sand tiger sharks arethreatened as bycatch of fisheries driven by abundant, wide-ranging scombroid fishes. The conservation status of someuncommon to rare deepwater lamnoids, including the goblin, megamouth and bigeye sand tiger sharks, is poorly known andis of concern as these sharks are found in areas subject to intensive deepwater and oceanic fisheries and are not beingmonitored.Local Names: Lamnoid sharks, Mackerel sharks.Remarks: Garman (1885) was the first author to propose a higher group name applicable to this order but the concept of anorder Lamniformes or equivalent ordinal taxon to include only lamnoid families is far more recent. Early writers and evensome modern authors generally grouped orectoloboids and sometimes carcharhinoids and other shark groups together withthe lamnoids. Garman’s Lamnae included most living sharks except hexanchoids, heterodontoids and squatinoids. Garman(1913) later recognized two primarily lamnoid ‘groups’ (equivalent to infraorders or superfamilies) in his suborder Antacea(sharks), the Carcharoidei (for odontaspidids and mitsukurinids), and the Isuroidei, for alopiids, lamnids, cetorhinids, and theorectoloboid whale shark, but did not place the two groups in a common higher lamnoid group. White (1936, 1937) andWhitley (1940) essentially followed Garman’s arrangement of dividing the lamnoids into two groups. An influential modernarrangement was Bigelow and Schroeder’s (1948) order Selachii, suborder Galeoidea, which included the lamnoid,orectoloboid, and carcharhinoid families in a single undifferentiated group. This has been followed by several authors invarious forms.Jordan (1923) was the first author to propose an exclusive group for the lamnoids, the series Lamnoidei (equivalent to aninfraorder or superfamily). Bertin’s (1939a) suborder Lamniformes included all lamnoids but also (inexplicably) thecarcharhinoid Pseudotriakis. Rass and Lindberg (1971) used the order Lamniformes exclusively for all lamnoids except thebasking shark, which was placed in the order Carcharhiniformes along with orectoloboids. Glikman (1967) proposed anorder Odontaspidida for all lamnoids except the basking shark, which was placed in the order Squatinida in a suborderSqualoidei including the squaloids. Compagno (1973) and Applegate (1974) reinvented the order Lamniformes in itsmodern form, essentially the same as Jordan’s series Lamnoidei, which has been followed by several authors includingCompagno (1984, 1988, 1990b, 1999), Cappetta (1987), Eschmeyer (1990, 1998), Nelson (1994), de Carvalho (1996), andShirai (1996). Chu and Meng (1979) used the order Isuriformes as an equivalent taxon to Lamniformes and Carroll (1988)used the order Galeomorpha, suborder Lamnoidei exclusively for lamnoids.Continuing work on the morphology of sharks by the writer as an extension of previous work (Compagno, 1990b) supportsthe retention of the Lamniformes as a monophyletic but morphologically and ecologically varied group. The arrangement oflamnoid families recognized in Compagno (1984, 1990b, 1999) are retained here, but the possibility remains that theOdontaspididae is paraphyletic, and that the two genera, Carcharias and Odontaspis, may be separable into two families,Carchariidae and Odontaspididae.Key to Families:1a. Snout greatly elongated and flattened, forming adagger-like blade; no precaudal pits; ventral caudallobe absent; anal fin broadly rounded(Fig. 42) . . . . . . . . . . . . family Mitsukurinidae1b. Snout very short to moderately elongated, conicalto flattened and broadly rounded but notblade-like; precaudal pits (upper pits, and oftenlowers) and ventral caudal lobe present; anal finangular (Fig. 43) . . . . . . . . . . . . . . . . . . 2Fig. 42 MitsukurinaFig. 43 Cetorhinus

54 FAO Species Catalogue for Fishery Purposes No. 12a. Snout very short and broadly rounded indorsoventral view; mouth terminal on head(Fig. 44); teeth in dental bands continuouslyvarying, no row groups; internal gill openingswith densely packed papillose gill rakers . . .. . . . . . . . . . . . . . . . . family Megachasmidae2b. Snout longer and narrowly to broadly parabolic indorsoventral view; mouth subterminal on head(Fig. 45); teeth differentiated into anteriors andlateroposteriors in upper jaw (Cetorhinidae), andanteriors, laterals and often intermediates andsymphysials in other taxa; internal gill openingseither without gill rakers or with rows of elongatedgill raker denticles (Cetorhinidae). . . . . . . . 3Fig. 44 Megachasma3a. Caudal fin about as long as rest of shark; last twogill openings above pectoral-fin base (Fig. 46). . . . . . . . . . . . . . . . . . . . . family Alopiidae3b. Caudal fin much shorter than rest of shark; all gillopenings in front of pectoral-fin base (Fig. 47) . . . . 4Fig. 45 Cetorhinus4a. Caudal fin asymmetrical, not lunate, ventral caudallobe short, preventral caudal margin muchshorter than dorsal caudal margin; caudalpeduncle without lateral keels or with weak ones(Fig. 47) . . . . . . . . . . . . . . . . . . . . . . . . 5Fig. 46 Alopias4b. Caudal fin nearly symmetrical and lunate, with along ventral lobe and preventral caudal marginnearly as long as dorsal caudal margin; caudalpeduncle with very strong lateral keels (Fig. 48) . . . . 65a. Eyes very large, body slender; anal fin narrow-based,pivoting; caudal peduncle with bothupper and lower precaudal pits and low lateralkeels on each side; gill openings extending ontodorsal surface of head (Fig. 49) . . . . . . . .. . . . . . . . . . . . . . . . family Pseudocarchariidae5b. Eyes relatively small, body stout; anal finbroad-based, not pivoting; caudal peduncle withan upper precaudal pit but without a lower pit orlateral keels; gill openings not extending ontodorsal surface of head (Fig. 47) . . family OdontaspididaeFig. 47 Carcharias6a. Teeth relatively few, enlarged and blade-like,with less than 40 rows in each jaw; gill openingslarge but ending far lateral to mid-dorsal surfaceof head (Fig. 48); internal gill openings withoutgill rakers . . . . . . . . . . . . . . . . . family Lamnidae6b. Teeth numerous, minute, hooked and notblade-like, with over 150 rows in each jaw; gillopenings extremely large, extending nearly tomid-dorsal surface of head (Fig. 45); internal gillopenings with prominent gill rakers formed frommodified dermal denticles . . . . . . . family CetorhinidaeFig. 48 IsurusFig. 49 Pseudocarcharias

Sharks of the World, Vol. 2 552.2.1 Family ODONTASPIDIDAEFamily: Family Odontaspides Müller and Henle, 1839, Syst. Beschr. Plagiost., pt. 2: 73. Emended to FamilyOdontaspididae Müller and Henle, 1839. The corrected form Odontaspididae was placed on the Official List of Family-GroupNames in Zoology (Name no. 385) but Odontaspides was placed on the Official Index of Rejected and Invalid Family-GroupNames in Zoology (Name no. 414) by the International Commission on Zoological Nomenclature (1965, Opinion 723, Bull.Zool. Nomencl., 22: 33, 34). Odontaspididae was given special endorsement by the International Commission on ZoologicalNomenclature (1987, Opinion 1459.6, Bull. Zool. Nomencl., 44(3): 216) to take precedence over Carchariidae Müller andHenle, 1838 when the two are synonymized.Type Genus: Odontaspis Agassiz, 1838.Number of Recognized Genera: 2.Synonyms: Subfamily Triglochidini Bonaparte, 1838: 208 (Family Squalidae). Type genus: Triglochis Müller and Henle,1837. Family Carchariae Müller and Henle, 1838d: 27. Type genus: Carcharias Rafinesque, 1810. Rejected by theInternational Commission on Zoological Nomenclature (1965, Opinion 723: 33) but reinstated by the Commission (1987,Opinion 1459.5: 216) in the corrected form Family Carchariidae Müller and Henle, 1838 on the Official List of Family-GroupNames in Zoology, with the special endorsement that it is not to be given precedence over Odontaspididae Müller andHenle, 1839 when considered a synonym of it. This name was widely used by earlier writers for members of theCarcharhinidae, following Müller and Henle’s original usage, but Jordan and Gilbert (1883: 27) and many subsequentwriters used it for members of the Odontaspididae with the assignment of Carcharias to this family. Family CarchariidaeJordan and Gilbert, 1883: 27. Emended spelling for Family Carchariae Müller and Henle, 1838. Type genus: CarchariasRafinesque, 1810. Family Eugomphodidae Applegate, Espinosa, Menchaca and Sotelo, 1979: 130. Type genus:Eugomphodus Gill, 1862. Also ibid.: 30, as Eugonphodidae, error for Eugomphodidae.FAO Names: En - Sand tiger sharks; Fr - Requins de sable; Sp - Solrayos, Toros.Field Marks: Large heavy-bodied sharks with conical to slightly depressed pointed snouts, long mouths extending behindeyes, small to moderately large eyes without nictitating eyelids, moderately long gill openings in front of pectoral origins,large teeth with slender cusps and lateral cusplets, small intermediate teeth separating anterior and lateral teeth in the upperjaw, two large dorsal fins and an anal fin, small pectoral fins, a compressed caudal peduncle without keels but with an upperprecaudal pit only, and an asymmetrical caudal fin with a strong but short ventral lobe.Diagnostic Features: Head much shorter than trunk. Snout short to moderately long, pointed and bulbously conical ormoderately depressed, not greatly elongated, flattened or blade-like. Eyes small to moderately large, length 1.4 to 4.1% ofprecaudal length. Gill openings moderately large, length of first 6.2 to 9.2% of precaudal length, not extending onto dorsalsurface of head; all gill openings anterior to pectoral-fin bases; no gill rakers on internal gill slits. Mouth large, parabolic,ventral on head; jaws strongly protrusable to almost opposite snout tip but not greatly distensible laterally. Teeth large,anteriors narrow and awl-like but laterals moderately compressed and blade-like, in 34 to 56/36 to 46 (71 to 102 total) rows,less than 60 rows in either jaw; 2 or 3 rows of large anterior teeth on each side of upper jaw, three rows in lower jaw, theuppers separated from the smaller upper lateral teeth by 1 to 5 rows of small intermediate teeth (rarely absent); one or morepairs of symphysial teeth present in the lower jaw or both jaws. Trunk compressed-cylindrical and moderately stout, firm andnot flabby. Caudal peduncle compressed and without keels but with a crescentic upper precaudal pit only. Dermal denticlesmoderately large and smooth, with flat crowns, small ridges and cusps, and with cusps directed posteriorly on lateraldenticles. Pectoral fins moderately long and broad, much shorter than head in adults; pectoral skeleton aplesodic withradials confined to fin bases. Pelvic fins large, nearly or quite as large as first dorsal fin; fin skeleton aplesodic. First dorsal finlarge, moderately high, erect and angular; first dorsal-fin skeleton aplesodic. Second dorsal and anal fins about as large asfirst dorsal fin or second dorsal smaller than first and as large or larger than anal fin; second dorsal and anal fins with broadnonpivoting bases. Caudal fin not lunate, dorsal lobe moderately long, less than half as long as rest of shark, ventral lobeshort but strong. Neurocranium low to moderately high, with a short to moderately elongated rostrum, depressed internasalseptum and widespread nasal capsules, small to large orbits with the supraorbital crests strong, small stapedial fenestrae,and with hyomandibular facets not extended outward. Vertebral centra strongly calcified with well-developed double conesand radii but no annuli. Total vertebral count 156 to 183, precaudal count 80 to 95, diplospondylous caudal count 71 to 88.Intestinal valve of ring type with 28 to 32 turns. Size large with adults 2.2 to at least 3.6 m.Distribution: Odontaspidids have a wide but sporadic geographic distribution in virtually all warm-temperate and tropicalseas, and further deepwater exploration and fisheries efforts with appropriate gear will undoubtedly reveal range extensions.Habitat: Sand tiger sharks are tropical to warm-temperate, inshore to offshore, littoral and deepwater sharks. They occur incontinental and insular waters from the outer shelves and down the slopes to possibly 1 600 m, on seamounts, and with onespecies (Odontaspis noronhai) also oceanic in the epipelagic and possibly the mesopelagic zone.Biology: Sand tiger sharks are relatively slow but active littoral, epibenthic and oceanic swimmers. They feed on a widevariety of bony fishes, other sharks, rays, squids and bottom crustaceans. Development is ovoviviparous (aplacentalviviparous), without a yolk-sac placenta but with uterine cannibalism in the genus Carcharias, in the form of both oophagyand adelphophagy (egg and embryo-eating).

56 FAO Species Catalogue for Fishery Purposes No. 1Interest to Fisheries and Human Impact: Sand tiger sharks, particularly Carcharias taurus and to a lesser extentOdontaspis ferox are or have been important for inshore and offshore fisheries wherever they occur, but are far lessimportant and less abundant than requiem sharks (Carcharhinidae) or hammerheads (Sphyrnidae). Odontaspis noronhaiis primarily a rare or uncommon bycatch of oceanic and slope line fisheries. In some areas of relative abundance, such asthe east coast of North America and Australia, C. taurus has severely declined due to fishing pressure (including attacks bydivers in Australia), but other areas that supported important fisheries (west Africa off Senegal, and the South China Sea)have not been monitored and local populations may need attention. Population trends in Odontaspis are essentiallyunknown, and potentially worrisome because of known catches by fisheries. Sand tiger sharks are presently protected in theeastern USA and Australia, and Carcharias taurus is to be decommercialized in South Africa and will only be fished bysports anglers.These sharks are inoffensive and usually not aggressive to humans in the water and are the subject of ecotouristic viewingby divers in South Africa, Australia, the east coast of the USA, the Mediterranean Sea, and Malpelo Island in the easternPacific. Swimmers, divers and fishermen commonly encounter (or formerly encountered) C. taurus and more recentlyOdontaspis ferox in areas of abundance, but despite their impressive teeth there have been few incidents of Carchariastaurus biting people, and none currently known for Odontaspis. As with other large sharks, sand tigers should be treatedwith respect and not harassed underwater.Carcharias taurus is important for aquarium displays worldwide, but Odontaspis species have not been kept in captivity tothe writer’s knowledge. The former has the ideal combination of a fearsome, showy, large ‘sharky’ appearance, combinedwith docility, hardiness, and great longevity in captivity.Local Names: Sand tiger sharks, Sand sharks, Ragged-tooth sharks, Grey or gray nurse sharks, Gray sharks, Patings,True sharks, Chuich’ih sha k’o; Mizuwani ka (Japan); Dlinnozubye akuly, Peschanye akuly (Russia); Tubaroes de areia.Remarks: The family Odontaspididae was recognized by many authors following Müller and Henle (1839). However,considerable confusion was caused by these authors (Müller and Henle, 1838d), who proposed Carchariidae forcarcharhinids but based it on Rafinesque’s (1810) genus Carcharias, which has as its type species the sand tiger shark C.taurus. Several authors followed Müller and Henle in using Carchariidae for carcharhinids and Odontaspididae (orOdontaspidae) for sand tiger sharks (Bleeker, 1859; Dumeril, 1865; Günther, 1870; Regan, 1906; Engelhardt, 1913), whileothers used Odontaspididae for sand tiger sharks and other names for carcharhinids (Gray, 1851; Gill, 1862b, 1872; Bertin,1939a; Berg, 1940; Berg and Svedovidov, 1955; Arambourg and Bertin, 1958; Norman, 1966). Carchariidae wasreestablished and widely used as a family for sand tiger sharks rather than carcharhinids (Jordan and Gilbert, 1883; Gill,1893; Jordan and Evermann, 1896; Garman, 1913; Jordan, 1923; Lozano y Rey, 1928; White, 1936, 1937; Whitley, 1940;Fowler, 1941, 1947; Bigelow and Schroeder, 1948; Matsubara, 1955; Garrick and Schultz, 1963; Romer, 1966; Pinchuk,1972; Chu and Meng, 1979; Carroll 1988). However, this was curtailed by White, Tucker and Marshall (1961), who proposedto validate the name Odontaspididae over Carchariidae due to greater use in the palaeontological literature. TheInternational Commission on Zoological Nomenclature (1965) suppressed Carchariidae for zoological literature. Variousauthors who published subsequent to this ruling have used Odontaspididae (Glikman, 1967; Patterson, 1967; Blot, 1969;Bailey et al, 1970; Budker and Whitehead, 1971; Lindberg, 1971; Rass and Lindberg, 1971; Compagno, 1973, 1981b, 1982,1984, 1999; Nelson, 1976, 1984, 1994; Gubanov, Kondyurin and Myagkov, 1986; Cappetta, 1987; Eschmeyer, 1990;Robins et al., 1991a; Shirai, 1996; Helfman, Collette and Facey, 1997; Eschmeyer, 1998). Applegate et al. (1979) usedEugomphodidae. However, following a petition by Compagno and Follett (1986), the International Commission onZoological Nomenclature (1987) reinstated Carchariidae as a valid family-group name for sand tiger sharks, but not havingprecedence over Odontaspididae unless removed from synonymy of that family. This leaves Carchariidae available for thegenus Carcharias and its fossil relatives if considered a distinct family from Odontaspididae and Odontaspis.Although most writers have recognized only one genus of Odontaspididae (or Carchariidae), two genera are recognizedhere for the living species. This follows recent palaeontological and neontological work (Glikman, 1964, 1967; Herman,1977; Compagno, 1981b, 1982, 1984; Cappetta, 1987) that treats the taurus and ferox groups of species as separategenera, but also morphological studies on the living species that revealed their distinctness (Compagno, 1984, 1990b;Compagno and Follett, 1987).The oldest genus-group names for the taurus group are Carcharias Rafinesque, 1810, and Triglochis Müller and Henle,1837, but these were rejected by the International Commission on Zoological Nomenclature (1965), following a proposal byWhite, Tucker and Marshall (1961). These authors reasoned that Carcharias Rafinesque, 1810 should be suppressedbecause the genus Odontaspis Agassiz, 1838 has been used far more frequently in the literature for odontaspidids(especially fossils) than Carcharias. A key point to their reasoning is that “...since the respective nominal type species ofCarcharias Rafinesque,1810, and Odontaspis J.L.R. Agassiz, 1838, are congeneric, it is the latter name which isthreatened by the former” (emphasis added). However, subsequent work indicated that the two type species, taurus andferox, were not congeneric, and, provided two genera of living odontaspidids are recognized, Odontaspis is not threatenedby Carcharias. Compagno (1977, 1981b, 1982, 1984), Welton and Zinsmeister (1980) and various subsequent writers usedthe genus Eugomphodus Gill, 1862 as the next valid name that could substitute for Carcharias. Cappetta (1987) usedSynodontaspis in favour of Eugomphodus, but the latter has priority. However, following the petition by Compagno and Follett(1986) the International Commission on Zoological Nomenclature (1987) reinstated the genus Carcharias Rafinesque, 1810.The genus Odontaspis as presently delimited is restricted to O. ferox, O. noronhai, and fossil species (Compagno, 1984;Cappetta, 1987), while the genus Carcharias has a single living species, C. taurus, and numerous fossil species.

Sharks of the World, Vol. 2 57Carcharias kamoharai Matsubara, 1936 (and its synonyms) have previously been placed in the genus Odontaspis orCarcharias (D’Aubrey, 1964a, b; Bass, D’Aubrey and Kistnasamy, 1975a), but this was placed in the genusPseudocarcharias and the family Pseudocarchariidae (Compagno, 1973, 1984).Phyletic studies by Compagno (1990b) based on external and skeletal morphology and dentition of living lamnoids, and agenetic study by Martin and Naylor (1997) suggested that Odontaspididae might be paraphyletic, with Carcharias andOdontaspis separate since the Cretaceous (Cappetta, 1987) and rating separate families. However, a phyletic study of thedentition of lamnoid sharks by Long and Waggoner (1996) suggested that Odontaspididae is monophyletic. Pending furtherstudy Odontaspis and Carcharias are retained in the family Odontaspididae following Compagno (1984).Literature: Garman (1913); Fowler (1941, 1967a); Bigelow and Schroeder (1948); Garrick and Schultz (1963); Lindberg(1971); Shiino (1972, 1976); Compagno (1973, 1982, 1984, 1990b, 1999); D’Aubrey (1964a, b); Bass, D’Aubrey andKistnasamy (1975a).Key to Genera:1a. Snout short and flattened; three rows of large upperanterior teeth on each side of symphysis; firstdorsal fin about as large or slightly larger thansecond dorsal fin and anal fin; first dorsal fincloser to pelvic-fin bases than pectoral-fin bases(Fig. 50) . . . . . . . . . . . . . . . . . . . . CarchariasFig. 50 Carcharias1b. Snout long and conical; two rows of large upperanterior teeth on each side of symphysis; firstdorsal fin noticeably larger than second dorsal finand anal fin; first dorsal fin closer to pectoral-finbases than pelvic-fin bases (Fig. 51) . . . . . OdontaspisFig. 51 OdontaspisCarcharias Rafinesque, 1810Genus: Carcharias Rafinesque, 1810, Caratt. gen. sp. anim. piant. Sicilia, Palermo, pt. 1: 10. Placed on the Official Index ofRejected and Invalid Generic Names in Zoology (Name no. 1746) by the International Commission on ZoologicalNomenclature (1965, Opinion 723.5a, Bull. Zool. Nomencl., 22(1): 33) following a proposal by White et al. (1961, Bull. Zool.Nomencl., 18(4): 277-278). However, Compagno and Follett (1986, Bull. Zool. Nomencl., 43(1): 89-92) argued for thereinstatement of Carcharias because its rejection on nomenclatural grounds interfered with taxonomic work on the family. This wasaccepted by the International Commission on Zoological Nomenclature with near-unanimity, and Carcharias was placed on theOfficial List of Generic Names in Zoology (1987, Opinion 1459.2, Bull. Zool. Nomencl., 44(3): 216), with the special endorsementthat it is not to be given precedence over Odontaspis Agassiz, 1838, whenever the two are considered synonyms.Type Species: Carcharias taurus Rafinesque, 1810, by monotypy (International Commission on Zoological Nomenclature,1912, Opinion 47, Smithsonian Pub., (2060): 108).Number of Recognized Species: 1.

58 FAO Species Catalogue for Fishery Purposes No. 1Synonyms: Genus Triglochis Müller and Henle, 1837a: 113. Placed on the Official Index of Rejected and Invalid GenericNames in Zoology (Name no. 1747) by the International Commission on Zoological Nomenclature (1965, Opinion 723.5b:33). Genus Eugomphodus Gill, 1862a: 60 (name only, but without allocated species); Gill, 1864: 260 (description). Typespecies, Eugomphodus griseus Gill, 1862, by monotypy, equals Carcharias griseus Storer, 1846 and C. griseus Ayres,1843, and a junior synonym of Carcharias taurus Rafinesque, 1810. Subgenus Synodontaspis White, 1931 (GenusOdontaspis Agassiz, 1838): 51. Type species, Carcharias taurus Rafinesque, 1810 by original designation. SubgenusWhite, 1931 (Genus Odontaspis Agassiz, 1838): 63. Type species, Odontaspis platensis Lahille, 1928, by originaldesignation.Diagnostic Features: Snout short with preoral length 0.3 to 0.5 times mouth width and 3.2 to 4.7% (usually 4.0 or less) oftotal length; snout somewhat flattened but not bulbously conical. Eyes smaller and about 0.9 to 1.4% of total length,dorsolateral in position with prominent lateral head ridges below them. Upper symphysial teeth usually absent; three rows oflarge upper anterior teeth on either side of symphysis; anterior teeth with stout broad-tipped cusps and short and stronglyhooked cusplets; anterior teeth enlarged with largest (second lower anterior) about 1.3 to 1.5% of total length; lateral teethcompressed, blade-like and with flattened cusps; posterior teeth strongly differentiated from lateral teeth, with cusps andcusplets reduced or absent and teeth molariform and carinate. Claspers tapering. First dorsal-fin base far posterior andcloser to pelvic-fin bases than to pectoral-fin bases, with pre-first dorsal-fin length 48 to 58% of precaudal length; firstdorsal-fin origin well behind inner margins of pectoral fins, insertion about over pelvic-fin origins. Second dorsal fin about aslarge as first dorsal fin. Anal fin about as large or slightly larger than dorsal fins; anal-fin origin under midbase of seconddorsal fin. Irises of eyes light greenish.Carcharias taurus Rafinesque, 1810 Fig. 52Carcharias taurus Rafinesque, 1810, Caratt. gen. sp. anim. piant. Sicilia, Palermo, pt. 1:10, pl. 14, fig. 1. Holotypeunknown; type locality, Sicily, Mediterranean Sea. Placed on the Official list of Specific Names in Zoology by theInternational Commission on Zoological Nomenclature (1987, Opinion 1459.4, Bull. Zool. Nomencl., 44(3): 216).Synonyms: Squalus americanus Mitchell, 1815: 483. No types known according to Eschmeyer (1998: CD-ROM). Typelocality, New York. Not Squalus americanus Gmelin, 1788, = Dalatias licha (Bonnaterre, 1788). Squalus macrodousMitchell, 1818: 328. Replacement name for S. americanus Mitchell, 1815. Squalus littoralis Le Sueur, 1818: 224. Holotype:91 cm specimen, New York, possibly not extant. Squalus littoralis Mitchell, 1818: 328. Types unknown. Apparently a juniorhomonym of S. littoralis Le Sueur, 1818 according to Eschmeyer (1998: CD-ROM). Carcharias griseus Ayres, 1842: 58-59(nomen nudum); Ayres, 1843a: 288 (no distinguishing features), Ayres, 1843b: 293, pl. 12, fig. 4. Type locality, Long Island,New York. No types according to Eschmeyer (1998: CD-ROM). Odontaspis americanus Abbott, 1861: 400 (newcombination validates name?). ?Carcharias tricuspidatus Day, 1878: 713, pl. 186, fig. 1. Figured from a skin about 373 cmTL, apparently lost (P.K. Talwar, pers. comm.), India (see remarks below). ?Odontaspis cinerea Macleay, in Ramsay, 1880:96. Port Jackson, New South Wales, name only, in footnote. ?Carcharias cuspidatus Ogilby, 1888?: 1767. Reference inFowler (1941: 122), but Ogilby, 1888 spelled it correctly as C. tricuspidatus, hence possibly an error by Fowler? Australia.Lamna ecarinata Hemprich and Ehrenberg, 1899: 8, pl. 6, fig. 1. Holotype, Zoologisches Museum, Museum für Naturkundeder Humboldt-Universität, Berlin, ZMB 4532, a stuffed 846 mm female according to Paepke and Schmidt (1988: 163),Alexandria, Mediterranean Sea. Carcharias arenarius Ogilby, 1911: 37. Holotype: Queensland Museum, Brisbane, QMI.1884 (dry mount) according to Eschmeyer (1998: CD-ROM), Moreton Bay, Queensland. Carcharias owstoni Garman,1913: 24. Holotype, Museum of Comparative Zoology, Harvard, MCZ-1278, 920 mm TL newborn or late foetal male, SagamiSea, Japan. Status confirmed by Hartel and Dingerkus (1997) with catalogue number MCZ-1278-S. ?Squalus lixaLarrañaga, 1923: 391. Reference from Eschmeyer (1998: CD-ROM); apparently a species dubium and nomen nudum,possibly referable to this species). Odontaspis platensis Lahille, 1928: 324, figs 13-16, pl. 3 (upper fig.). Syntypes: Possiblythree syntypes, a 2.33 m specimen from Panela, near Montevideo, Uruguay, a jaw from Necochea, Quequen, Argentina,and a 2.27 m female (on which the description is primarily based) from Bahia Blanca, Argentina, status uncertain.Odontaspis tricuspitatus Fang and Wang, 1932: 241, fig. 12. Chefoo, China, apparently a spelling error for Carchariastricuspidatus.Other Combinations: Eugomphodus taurus (Rafinesque, 1810), Odontaspis taurus (Rafinesque, 1810), Triglochistaurus (Rafinesque, 1810), Synodontaspis taurus (Rafinesque, 1810), Carcharias littoralis (Le Sueur, 1818), Odontaspisgriseus (Ayres, 1843), Eugomphodus griseus (Ayres, 1843), Odontaspis littoralis (Le Sueur, 1818), Eugomphoduslittoralis (Le Sueur, 1818), Carcharias americanus (Mitchell, 1815), Odontaspis tricuspidatus (Day, 1878), Carchariasplatensis (Lahille, 1928).

Sharks of the World, Vol. 2 59FAO Names: En - Sand tiger shark; Fr - Requin taureau; Sp - Toro bacota.UPPER AND LOWER TEETH OF LEFT SIDEUNDERSIDE OF HEADField Marks: A large, bulky shark with a flattened-conical snout, eyes without nictitating eyelids, mouth long and extendingbehind eyes, teeth large with prominent narrow cusps and lateral cusplets, upper anterior teeth separated from lateral teethby small intermediate teeth, anal fin and both dorsal fins equally large and broad-based, first dorsal fin on back closer topelvic fins than to pectoral fins, upper precaudal pit present but lateral keels absent from caudal peduncle, caudal finasymmetrical but with a strong ventral lobe. Colour: light brown, often with darker reddish or brownish spots scattered onbody, eyes with light green irises.Diagnostic Features: See genus Carcharias above.Fig. 52 Carcharias taurusDistribution: Wide-ranging in warm-temperate and tropical coastal waters of the Atlantic Ocean, Mediterranean Sea, andIndo-West Pacific Ocean; absent from the Central Pacific and eastern Pacific Oceans. Western Atlantic: Canada (Gulf ofMaine: New Brunswick, rare), USA (Cape Cod, Massachusetts, New York, Rhode Island, Connecticut, New Jersey,Delaware, Maryland, Virginia, North Carolina, South Carolina to Florida, northern Gulf of Mexico including western Florida,Louisiana and Texas), Bahamas, Bermuda; southern Brazil to Uruguay and Argentina. Eastern Atlantic: Mediterranean toCanary Islands, Morocco, Western Sahara, Mauritania, Sao Tome and Principe, Cape Verde Islands, Senegal, Ghana,southern Nigeria to Cameroon, Angola, Namibia, and South Africa (Western Cape). Indo-West Pacific: South Africa (entireeast coast), Mozambique, possibly Providence, Farquhar and Cerf Islands (western Indian Ocean), Red Sea, ?Oman,Pakistan, ?India. Western Pacific: ?Indonesia (Laiwui, Obi Major, Aru Islands), ?Malaysia, Viet Nam, ?Philippines, Japan,Taiwan (Province of China), China; Australia (Queensland, New South Wales, Victoria, South Australia, Northern Territory,Western Australia; Tasmanian records need confirmation).

60 FAO Species Catalogue for Fishery Purposes No. 1Habitat: An inshore and offshore, littoral shark. This shark occurs in the surf zone off sandy and rocky beaches, in shallowbays, on offshore banks and reefs, in underwater caves, in troughs on sandy areas, and around coral and rocky reefs fromthe intertidal less than 1 m deep down to at least 191 m, with most at depths of 15 to 25 m. This species is often found near oron the bottom but also occurs in midwater or at the surface.Biology: A common or formerly common to abundant shark, present in large aggregations during breeding. It is a strong butslow midwater swimmer that is more active at night. This shark is denser than water, but it swallows air at the surface andholds it in its stomach to maintain approximately neutral buoyancy. Like a bony fish with a swim bladder, it can readily haltand hover motionless in the water (stalling).This species occurs as solitary individuals or in small to large aggregations or schools, ranging from 20 to at least 80individuals in Australia and South Africa. It is strongly migratory in parts of its range, particularly in its northern and southernextremities where pronounced poleward migrations occur in summer and equatorial movements in autumn and winter(Australia, the east coast of the USA, and the east coast of South Africa). Aggregations of individuals occur for feeding,courtship, mating and birth. In Australia sexual segregation and separate migration of the sexes may occur off the east coastof Australia, with males predominant off southern Queensland during the winter and females off New South Wales. Off SouthAfrica courtship and mating apparently occurs in the more tropical parts of its range, while pregnant females give birth inwarm-temperate waters. In North America, nursery areas are not well-defined, and it may give birth along its entire rangethere during March and April.The behaviour of this shark has been studied in captivity (Gordon, 1993), in a large shark tank housing sand tigers (youngand adult), wobbegongs (Orectolobus), grey sharks (Carcharhinus), sevengills (Notorynchus), stingrays, and reef andpelagic teleosts. These studies have shown evidence of complex behaviour, including social interactions (with courtship andmating) between three adult sharks and asocial interactions with other marine vertebrates and divers. A dominancehierarchy was recorded among three adult sharks (two males and a female). Behaviour patterns observed include stalling(shark stops swimming and hovers above the bottom), nosing (male comes behind and below female, and places his snoutjust below her cloaca); tailing (male closely follows another male, his head interfering with normal caudal movement of theleading shark), submissive behaviour (or bowing, female swims slowly, with body angled about 15° downward, exposing thepelvic fins, and just prior to copulation), cupping and flaring (female depresses her pelvic fins into a cup-shape, then flaresthem outward, exposing the cloaca), snapping (males give quick bites to other fishes in the tank and then withdraws),stalking (close circling and passing of other fishes and divers), clasper flexion (movements of individual claspers forwardand rearward), clasper splaying (claspers spread laterally), clasper crossing (claspers crossed with tips posterolateral),shielding (female swims close to bottom to protect her cloaca), and scarring (infliction of light bites by both males andfemales).Most of the behaviours were observed during courtship and mating. The two adult males became reluctant to feed, andaggressive toward other fishes, particularly the grey sharks. The adult female slowed over large sand flats, cupped herpelvic fins when a male approached, and the two males lightened in colour and began ‘defending’ the sand flat area andpresumably the female, snapping, stalking and tailing at immature sand tigers, other sharks and divers. Snapping and tailingresulted in one male driving off the other without scarring it. The dominant male then would bite the female on the anal finforward to the pectoral fin, and the female would turn and bite the male, then return to patrol the sandy areas. Shielding alsooccurred, and cupping and flaring of the female’s pelvic fins, while the male responded by nosing. Over a few days ofcourtship between the dominant male and the female, the dominant male eventually grabbed the female by the right flankand pectoral fin, contorted his body toward the female, and inserted his right clasper in her cloaca for a few minutes, thenbroke away and showed little further interest. Additional scarring on the female suggested that a secondary mating hadoccurred in one instance.Reproduction in this species is better known than in most other lamnoids and features uterine cannibalism or cannibalvivipary. Gilmore, Dodrill and Linley (1983) give a detailed and fascinating account of the reproduction of the sand tigershark, documenting the sequence of intrauterine nourishment in this species, in which a successful embryo progresses fromusing stored yolk through killing and eating other embryos to eating unfertilized eggs, for a 9 to 12-month gestation period.There are normally two young in a litter, one per uterus. Eggs leave the ovaries, and while in transit in the oviducts arefertilized and enclosed in groups of 16 to 23 in egg cases. However, at some time between fertilization and birth only oneembryo of its group prevails, apparently by devouring its rivals, and this proceeds to eat fertilized eggs and smaller potentialsiblings in utero until birth. Unlike ovoviviparous non-cannibal and viviparous species, the yolk sac is reabsorbed at a smallsize, less than 17 cm, and the umbilical scar may be lost. At 17 cm, foetuses have sharp, functional teeth and are feeding; atabout 26 cm, they can swim in utero; late foetuses are near 1 m long and one bit the hand of an investigating scientist(Stewart Springer). This species may breed every other year, with a rest year between pregnancies.This species is thought to deposit two growth rings in its vertebrae per year, with males becoming adult at about five yearsand females at over six. Growth rings indicate animals growing to at least ten years old, with one adult male surviving nearly17 years in captivity in South Africa. Growth is rapid over the first five years, but declines beyond 10 years and is virtuallyabsent at 16 years.The sand tiger primarily feeds on a wide variety of bony fishes (teleosts), with elasmobranchs an important secondary prey;crustaceans, cephalopods, and marine mammals are also taken. Bony fish prey includes herring (Clupeidae), anchovies(Engraulidae), hake (Merluccidae), eels (Anguillidae), monkfish or anglers (Lophiidae), cusk eels (Ophidiidae), lizardfish(Synodontidae), sea catfish (Ariidae), croakers (Sciaenidae), Australian salmon (Arripidae), morwong (Cheilodactylidae),

Sharks of the World, Vol. 2 61rock blackfish or opaleyes (Girellidae), bluefish, elf or taylor (Pomatomidae), mackerel and bonito (Scombridae),butterfishes (Stromateidae), snappers (Lutjanidae), wrasses (Labridae), mullet (Mugilidae), spadefish (Chaetodipteridae),sea robins (Triglidae), flatheads (Platycephalidae), duckbills (Percophidae), midshipmen (Batrachodidae), sea basses(Serranidae), porgies or sea bream (Sparidae), jacks (Carangidae), remoras (Echeneidae), flatfish (Pleuronectiformes)including soles (Soleidae), American soles (Achiridae), Atlantic flounders (Scophthalmidae), and righteye flounders(Paralichthyidae), and undoubtedly many others. Elasmobranch prey includes requiem sharks (Carcharhinidae),houndsharks (Triakidae), angel sharks (Squatinidae), skates (Rajidae) and their egg cases, and eagle rays (Aetobatus andMyliobatis, Myliobatidae). Invertebrate prey includes squid (Loliginidae), crabs, lobsters and hermit crabs (Paguridae).Plant material is rarely found in stomach contents, and presumably is accidentally ingested along with animal prey. OffUruguay, Praderi (1985) found pinniped remains (7 individuals out of 12 identified as sea lion, Otaria) in 2% of 557 stomachsof the sand tiger shark, as well as wounds attributable to this species on two Franciscana dolphins (Pontoporia blainvillei;not necessarily from predation). Schools of this shark have been observed feeding cooperatively, surrounding and bunchingschooling prey and then feeding on them, and may use tail-slapping to scare and confuse schooling carangids. Australianand South African sharks sometimes have hydroid growths on their teeth, suggesting that the sharks were not feeding at thetime.Size: Maximum total length at least 318 cm, with recent records from shark meshing operations in Australia suggesting amaximum of at least 4.3 m. Older accounts attribute a size of at least 6.1 m to this species (Day, 1878), but this is unlikely.Size at birth 95 to 105 cm. Males maturing at about 190 to 195 cm with adults 220 to 257+ cm; females maturing at 220 cm ormore and reaching 300+ cm, with immatures up to 225 cm.Interest to Fisheries and Human Impact: This shark is generally fished commercially wherever it occurs, but particularly inthe western North Pacific (where it is highly prized for food), off tropical west Africa, in the northern Indian Ocean (India andPakistan), and in the western Atlantic (where it has a lesser value as a food-fish in the western North Atlantic but has beenfished for hides and fins). Caught with line gear, but also with fixed bottom gill nets and in pelagic and bottom trawls. It is ofonly moderate importance to commercial fisheries in comparison to more abundant carcharhinids. The meat of this shark isutilized for human consumption (utilized fresh, frozen, smoked and dried-salted), carcasses for fishmeal, hides for leather(the skin is very thick and tough, but hides can be spoiled by mating scars), liver for oil, fins for the oriental sharkfin trade, andjaws and teeth for trophies and ornaments. In Australia (New South Wales) the flesh has been found to have high mercurylevels (about ten times the permitted maximum), making the species unsuitable for human consumption.The sand tiger is sought by anglers in fishing competitions because of its size (South Africa), but it is sluggish when hookedand generally does not give much of a fight. Divers using powerheads and poisoned spears found this slow-moving speciesan easy target in Australia and to a much more limited extent in South Africa. Such crude and barbaric sport, analogous toshooting domestic cattle with a pistol, caused a severe decline in the number of these sharks in Australia and has beenbanned since 1984. South African divers stopped shooting ragged-tooth sharks many years ago. Australian anglers agreedto a ban on catching sand tiger sharks in 1979. These sharks are caught live for the public aquarium trade, but the number ofsharks involved in the trade is not recorded and is probably small. Separate catch statistics for this shark are not reported toFAO.This species formerly had a bad reputation as a ‘maneater’ in Australian waters, but apart from shark-attack hyperbole this isapparently due to confusion with other species, particularly requiem sharks (Carcharhinidae) but possibly the white shark.Observations of this shark underwater suggest that it is primarily inoffensive and not aggressive toward people when notprovoked, though its size and jagged dentition should invite respect. However, it occasionally harasses and rarely bitesdivers, particularly when they are spearfishing or possibly when adults are courting. This shark will steal fish off stringers andspears underwater, underlining the desirability of boating one’s catch when this shark or others are about. Relatively fewvalid instances of this shark biting people have been reported, and minimalist bites on divers off South Africa and elsewheresuggest non-predatory behaviour and possibly agonistic activity similar to scarring bites delivered to conspecifics, otherspecies of sharks and occasionally to divers in aquaria. Divers view aggregations of these sharks off Australia (particularlyoff New South Wales), South Africa and the east coast of North America. There are regular charter tours that include placesfrequented by congregations of these sharks. Interest in ecotouristic diving on sand tiger sharks is high in Australia, leadingto potential problems such as too many divers driving away sharks in a given area, and feeding of sharks resulting inpossible incidents of biting. Management of diving contact with sand tiger sharks may be required in the future, as with whitesharks. Capture, tagging and live release of sand tiger sharks by recreational anglers and scientists occurs in South Africaand elsewhere, and sonic tagging is planned in Australia to trace movements.This shark is prized as an exhibit in large aquaria and oceanaria and is regularly shown in the United States, Europe,Australia and South Africa. It is very hardy and docile in captivity, grows to adult size and can live for decades in aquaria, canbe kept with other sharks and fishes (although it sometimes may prey on fishes and will bite and sometimes kill othersharks), and makes an impressive display because of its showy, toothy, ‘sharky’ appearance, large size, and amenability tobeing approached and even fed underwater by divers. An aquarium in Cape Town, South Africa allows ecotouristic divers toswim with these sharks in their giant oceanic tank. Adults will court and even give birth in adequate tank facilities.The conservation status of this species is of concern because of its vulnerability to fisheries and recorded declines offeastern North America and Australia. It is listed as a vulnerable species by IUCN in its Red List of Threatened Animals. Itcongregates in large numbers in coastal areas during the breeding season, and is readily accessible to commercial fisherswho can catch it with little effort. It began to decline dramatically off New South Wales, Australia, in the 1960s due tofisheries, including shooting by sports divers, and catches in anti-shark nets dropped thirty-fold. It was fully protected off

62 FAO Species Catalogue for Fishery Purposes No. 1New South Wales in 1984, and later off Queensland and in all Commonwealth waters of Australia, as well as in federalwaters of the eastern coast of the USA. However, despite it having a midrange intrinsic rebound potential (Smith, Au andShow, 1998), the population off New South Wales has not shown signs of recovery and a new recovery plan is being draftedby the Australian Government (Environment Australia, 2000a).A burgeoning fishery for sand tigers along the east coast of the United States during the 1990s caused local catches toplummet off North Carolina, Florida and Chesapeake Bay, and the species was given total protection from sports andcommercial fisheries in 1997.There is a proposal under consideration by the South African Sea Fisheries Research Institute (now Marine and CoastalManagement) to decommercialize catches of this species, allowing it only to be fished by sports anglers. Unfortunately,relatively little is known of trends in other places in which sand tiger sharks have been heavily fished, including the westernNorth Pacific, western South Atlantic and eastern North Atlantic.Local Names: Sand shark, Sand tiger shark, Sand tiger (USA); Tiburón, Sarda, Tiburón de leznas (Argentina); Lamio,Verdoun, Odontaspide taureau (France); Odontaspe tauro, Carcharia tauro, Triglochide tauro, Pisci cani (Italy); Pez toro(Spain); Psina zmijozuba sivka (Adriatic); Sarda (Canaries); Requin sable, Requin sable tachete (Senegal); Grey nurseshark, Grey nurse, Sand shark, Shovel-nosed shark (Australia); Shirowani, Umiwani, Owston’s sand shark (Japan);Shovelnose shark, Sand tiger, Yellow shark, Brown shark, Grey shark, Tiger shark, Grey-nurse (English); Ragged-tooth orRaggedtooth shark, Spotted ragged-tooth or Raggedtooth shark, Raggie, Yellow belly, Blue nurse shark,Spikkel-skeurtandhaai (South Africa); Indian sand tiger, Requin taureau bambak, Toro bambaco, Blue nurse shark, Bluenurse, Dundanee (India); Ca nham nhon, Ca nham nhan (Viet Nam).Remarks: Included as synonyms of Carcharias taurus are a number of regional species that have often been consideredvalid in the older literature, but which are most likely local representatives (or at most local subspecies) of a single,wide-ranging species. The dentitional characters most often used to distinguish several of these species (see Bigelow andSchroeder, 1948) apparently vary considerably within samples from a given area (Applegate, 1965; Sadowsky, 1970;Taniuchi, 1970). Regional names include Squalus americanus and its synonyms from the western North Atlantic,Odontaspis platensis from the western South Atlantic, Carcharias tricuspidatus from the Indian subcontinent, C. owstonifrom the western North Pacific, and C. arenarius from Australia. Abe et al. (1968, 1969), Sadowsky (1970), Taniuchi (1970),and Whitley and Pollard (1980) have all used the species name taurus for the local representatives of the species formerlynamed owstoni, arenarius and platensis. Carcharias taurus itself has been placed in different genera as Odontaspistaurus, Eugomphodus taurus and Synodontaspis taurus.Carcharias tricuspidatus was distinguished by Compagno (1984) from C. taurus by characters from Day’s (1878) originaldescription. A problem was that these characters could not be confirmed in recent accounts or specimens of this speciesfrom the Indian subcontinent. Compagno (1984) suggested that C. tricuspidatus may be a junior synonym of C. taurus aswith other regional species of Carcharias. Day’s (1878) original description stated that C. tricuspidatus lacked labialfurrows and pictured it as having a broadly rounded snout (C. taurus has a rounded-angular snout and well-developed labialfurrows). However, since the species was described from a skin, now lost, it is possible that the reported snout shape is inerror and that the labial furrows were overlooked. Day’s (1878, pl. 186, fig. 13) drawing of the snout of his C. tricuspidatusmay even be based on Negaprion acutidens, though the lateral view and tooth illustrated are very similar to those ofCarcharias taurus. An alternative is that the drawing is inaccurate and sketchy, as with several others of Day’s sharkdrawings, and might reflect problems Day had in interpreting the holotype. During a trip to India in 1982, the writer wasunable to examine any specimens of Carcharias from India (which is apparently much less common there than largecarcharhinids) and was unable to confirm or deny the validity of this species. Compagno (1984) left this as a species dubium,but further consideration leads me to place it as a tentative synonym of C. taurus as suggested by C. taurus material Iexamined in the Pakistan collection of the Los Angeles County Museum of Natural History.Literature: Rafinesque (1810); Day (1878); Garman (1913); Whitley (1940); Bigelow and Schroeder (1948); Springer(1948); Cadenat (1956); D’Aubrey (1964a,b); Applegate (1965); Abe et al. (1968, 1969); Lineaweaver and Backus (1970);Taniuchi (1970); Sadowsky (1970); Bass, D’Aubrey and Kistnasamy (1975a); Gilmore, Dodril and Linley (1983); Compagno(1984); Govender, Kistnasamy and van der Elst (1991); Gordon (1993); Michael (1993); Musick, Branstetter andColvocoresses (1993); Branstetter and Musick (1994); Last and Stevens (1994); Pollard, Smith and Smith (1996); Hoeseand Moore (1998); McEachran and Fechhelm (1998); Smith, Au and Show (1998); Castro, Woodley and Brudek (1999);Gelsleichter, Musick and Nichols (1999); Environment Australia (2000a).

Sharks of the World, Vol. 2 63Odontaspis Agassiz, 1838Genus: Odontaspis Agassiz, 1838, Recher. Poiss. Foss., 3: 86, 87. Placed on the Official List of Generic Names in Zoology(Name no. 1659) by the International Commission on Zoological Nomenclature (1965, Opinion 723.3c, Bull. Zool. Nomencl.,22(1): 33).Type Species: Carcharias ferox Risso, 1826 by monotypy, equals Squalus ferox Risso, 1810. This genus takesprecedence over Carcharias Rafinesque, 1810 when the two are considered synonyms, by special endorsement of theInternational Commission on Zoological Nomenclature (1987, Opinion 1459.3, Bull. Zool. Nomencl., 44(3): 216).Number of Recognized Species: 2.Synonyms: None.Field Marks: Large bulky sharks with bulbous conical snouts, eyes without nictitating eyelids, mouth long and extendingbehind eyes, teeth moderately large with prominent narrow cusps and lateral cusplets, upper anterior teeth separated fromlateral teeth by small intermediate teeth, anal fin and second dorsal fin smaller than first dorsal fin, all three broad-based, firstdorsal fin on back closer to pectoral fins than to pelvic fins, only upper precaudal pit present but lateral keels absent fromcaudal peduncle, caudal fin asymmetrical but with a strong ventral lobe. Colour: blackish to light brown, eyes with blackirises.Diagnostic Features: Snout longer with preoral length 0.8 to 1.2 times mouth width and 4.4 to 7.8% (usually over 5%) oftotal length; snout long and bulbously conical. Eyes moderately large and about 1.6 to 2.8% of total length, lateral in positionwithout lateral-head ridges below them. A pair of upper symphysial tooth rows present or absent; two rows of large upperanterior teeth on either side of symphysis; anterior teeth with slender narrow-tipped cusps and long straight or weakly curvedbut not hooked cusplets; anterior teeth smaller with largest (second lower anterior) about 1% of total length; lateral teeth littlecompressed and not blade-like and with little-flattened cusps; posterior teeth grading into lateral teeth, with prominent cuspsand cusplets and not molariform. Claspers cylindrical and blunt-tipped. First dorsal-fin base more anterior and closer topectoral-fin bases than to pelvic-fin bases and with pre-first dorsal-fin space 43 to 49% of precaudal length; first dorsal-finorigin over inner margins of pectoral fins, insertion well anterior to pelvic-fin origins. Second dorsal fin about half as large asfirst dorsal fin. Anal fin subequal to or somewhat smaller than second dorsal; anal-fin origin under or behind seconddorsal-fin insertion. Irises of eyes black.Local Names: Deepwater sand tigers.Remarks: Following Compagno (1984) and Cappetta (1987), this account restricts the genus Odontaspis to species relatedto O. ferox, including the living O. ferox, O. noronhai, and fossil species. O. noronhai needs further study of its anatomy toclarify differences between it and O. ferox and between Odontaspis and Carcharias. Data on vertebral counts, vertebralcalcification patterns, cranial and fin skeletal morphology, and clasper morphology were unavailable for Odontaspisnoronhai.Key to Species:1a. Teeth mostly with two or threecusplets on each side of cusp(Fig. 53); colour grey or grey-brownabove, lighter below, often with darker2-3 cuspletsspots on sides but without awhite-tipped first dorsal fin . . . . Odontaspis ferox1 cusplet1b. Teeth mostly with only one cusplet oneach side of cusp (Fig. 54); colourdark reddish brown to blackish brownor black above and below, first dorsalfin often with a white blotch on its tip. . . . . . . . . . . . . . . . Odontaspis noronhaiUPPER ANTERIOR TOOTHFig. 53 Odontaspis feroxUPPER ANTERIOR TOOTHFig. 54 Odontaspis noronhai

64 FAO Species Catalogue for Fishery Purposes No. 1Odontaspis ferox (Risso, 1810) Fig. 55Squalus ferox Risso, 1810, Ichthyol. Nice, Paris: 38. Holotype unknown; type locality off Nice, France, in the MediterraneanSea. Also, Carcharias ferox Risso, 1826, Hist. nat. Princip. Prod. Europe Méred., Paris, Poissons, 3: 122. Descriptionvirtually verbatim that of Squalus ferox Risso, 1810, and quite evidently a generic translocation, not a new species name.Placed on the Official List of Specific Names in Zoology (Name no. 2057) by the International Commission on ZoologicalNomenclature (1965, Opinion 723.4.c: 33).Synonyms: Odontaspis herbsti Whitley, 1950: 234, fig. 1, pl. 17, fig. 1. Holotype: Australian Museum, Sydney,AMS-IB.2136, 168 cm immature male, Gabo Island, New South Wales, 137 m depth.Other Combinations: None.FAO Names: En - Smalltooth sandtiger; Fr - Requin féroce; Sp - Solrayo.UPPER AND LOWER TEETH ON LEFT SIDEFig. 55 Odontaspis feroxUNDERSIDE OF HEADField Marks: A large, bulky shark with a long bulbously conical snout, eyes moderately large without nictitating eyelids,mouth long and extending behind eyes, teeth moderately large with prominent narrow cusps and two or more pairs of lateralcusplets, upper anterior teeth separated from lateral teeth by 2 to 5 rows of small intermediate teeth, anal fin and seconddorsal fin smaller than first dorsal fin but broad-based, first dorsal fin on back and closer to pectoral fins than pelvic fins,upper precaudal pit present but lateral keels absent from caudal peduncle, caudal fin asymmetrical but with a strong ventrallobe, colour medium grey or grey-brown above, usually lighter below, sometimes with darker spots scattered on body.Diagnostic Features: Two to five (mostly four) rows of small intermediate teeth between upper anterior and lateral toothrows; a pair of upper and a pair of lower symphysial teeth present; tooth rows numerous, 48 to 56/36 to 46 (88 to 102 total);root lobes of anterolateral teeth deeply arched and narrow; anterolateral teeth usually with 2 or 3 pairs of lateral cusplets.Pectoral fins angular. First dorsal apex subangular in adults. Anal fin with height 4.6 to 6.0% of total length. Anal fin withstrongly concave posterior margin. Caudal fin with ventral caudal lobe short but stout. Colour: medium grey or grey-brownabove, lighter below, with darker dusky spots on sides of some individuals; fins dusky with blackish edges in young butuniform dusky in adults, first dorsal fin without a white blotch.Distribution: Possibly circumglobal in warm-temperate and tropical waters but spottily distributed. Western Atlantic:Mexico (Campeche Bank), United States (North Carolina), Brazil (NE Natal). Eastern North Atlantic: France (Bay of Biscay),Madeira, Morocco, Western Sahara, Mediterranean (Algeria, Italy, Adriatic, Lebanon). Western Indian Ocean: South Africa(KwaZulu-Natal), NE and SE of Madagascar in open ocean?, Maldives, open ocean SSE of Sri Lanka, SW of Sumatra andW of northern Australia. Western Pacific: Japan (Oshina Islands, SE Honshu off Izu-Shichito Islands), Australia (New SouthWales), New Zealand and Kermadec Islands. Central Pacific: Hawaiian Islands (Oahu; record of “O. owstoni” from PedestalSeamount in the Hawaiian range possibly this species or O. noronhai?). Eastern Pacific: United States (southernCalifornia), Mexico (Gulf of California, Baja California), Colombia (Malpelo Island). A cosmopolitan distribution has beenproposed for this species (Bonfil, 1995) and further exploration of deepish waters around the world might unveil its presencein still unknown parts of its range.

Sharks of the World, Vol. 2 65Habitat: This shark is a little-known inhabitant of deepish water in warm-temperate and tropical seas, on or near the bottomon continental and insular shelves and upper slopes at depths of 13 to 420 m, also possibly epipelagic zone in 140 to 180 mover the ocean floor. Sometimes observed by divers near dropoffs on coral reefs.Biology: An active-swimming offshore shark, caught and seen as individuals and in small groups. Reproduction is sketchilyknown in the species, with litter size unknown. An adult female from the Gulf of California had its right ovary filled withhundreds of small eggs and suggests that the species practices uterine cannibalism in the form of oophagy. It is not known ifdeveloping foetuses attack and devour each other until only one is left in each uterus as in Carcharias taurus, or if largerlitters are possible.This species feeds on small bony fishes, squid and shrimp. The teeth of this species and of Odontaspis noronhai arenoticeably smaller and less robust than those of Carcharias taurus (with this species having larger anterior teeth thanOdontaspis noronhai), suggesting that both species take smaller and possibly less active prey than Carcharias taurus.Also, the dentitions of both species are more weakly differentiated along the jaws, with their lateral teeth less specialized forcutting than Carcharias taurus and their posterior teeth not differentiated into specialized crushers. This suggests a moreuniform diet of softer prey than in C. taurus.Apparently the offshore and deepish-water habitat of this species does not allow this shark to regulate its buoyancy bygulping air as in C. taurus; it does however, have a longer body cavity than C. taurus, with a very large, oily liver, andpresumably uses this organ as its primary hydrostatic structure.Off Lebanon, adult individuals confronted by scuba divers may approach quite closely or slowly flee, and have been seen tohover (stall), gape,doaU-turn and do tail-shake (shaking their caudal fins vigorously) which may indicate ambiguity or mildagonistic reactions to divers or possibly conspecifics. Individuals have been seen with scars possibly from courtship orpossibly low-intensity scarring. They occur singly and in small groups and seem to prefer swimming near the bottom, insandy areas and on rocky reefs, sometimes following gullies or depressions in the reef (I.K. Fergusson, L.J.V. Compagno,K.J. Graham, F. Fakhoury, W. Noshie and W. Noshie, unpublished data).Size: Maximum total length at least 410 cm and possibly larger; size at birth above 105 cm; male mature at 275 cm, femalesadult at 364 cm. Specimens recently seen by divers at Malpelo Island are said to be considerably larger than the knownmaximum, but specimens have not been measured or photographed in such a way as to confirm this.Interest to Fisheries and Human Impact: This uncommon to rare but wide-ranging species is primarily fished in theMediterranean Sea and off Japan with bottom gill nets, line gear, and bottom trawls, and less commonly elsewhere. It formsa discarded or utilized bycatch of deepwater line and net fisheries in areas where it occurs. It is used in Japan for humanconsumption and for its liver, which is very large and oily, and has a reasonably high squalene content. Its flesh is consideredfar inferior to that of Carcharias taurus in Japan. This shark has not been recorded as biting people, and recent underwaterobservations by ecotouristic divers in the Mediterranean and off Malpelo Island in the eastern Pacific suggest that it isessentially docile although sometimes inquisitive when confronted by people. Its conservation status is essentiallyunknown. It has been protected in Australia since 1984.

66 FAO Species Catalogue for Fishery Purposes No. 1Local Names: Bumpytail ragged-tooth shark, Bumpytail ragged-tooth, Ragged-tooth, Knopstert-skeurtandhaai (SouthAfrica); Sand tiger shark, Herbsts or Herbst’s nurse shark, Smalltooth sand tiger (Australia); Tiger ragged-tooth (MalpeloIsland); Cagnia, Can da denti, Cagnassown de foundo, Carcaria feroce, Lamia, Odontaspe feroce, Pisci cani, Smidiru,Squalo feroce, Triglochide feroce (Italy); Salroig, Surraig, Solraig (Spain); Smalltooth sand tiger, Ragged-tooth shark(Azores); Psina zmij ozuba ruzicua (Adriatic); Ragged-tooth shark (USA, California), Fierce shark, Ôwanizame (Japan).Remarks: Garrick (1974) recognized Odontaspis herbsti for members of the genus from New Zealand, Australia, Californiaand Madeira that differed from the Mediterranean O. ferox only in lacking spots. This was followed by Bass, D’Aubrey andKistnasamy (1975a) for South African spotless individuals, but Robins et al. (1980: 69) note that specimens from Californiamay have spots or lack them. Observations of live O. ferox underwater in the Mediterranean Sea and off Malpelo Island alsoreveals individuals with and without spots (I.K. Fergusson, L.J.V. Compagno. K.J. Graham, F. Fakhoury, W. Noshie andW. Noshie, unpublished data). Apparently presence of spots reflects individual variation in a single species. Carchariastaurus is also variable in presence or absence of spots.Literature: Risso (1810); Garman (1913); Maul (1955); Tortonese (1956); Daugherty (1964); D’Aubrey (1964a, b); Abe et al.(1968); Garrick (1974); Bass, D’Aubrey and Kistnasamy (1975); Taniuchi (1983); Compagno (1984); Nakaya (1984); Quero(1984); Gubanov (1985); Seigel and Compagno (1986); Springer (1990); Anderson and Ahmed (1993); Michael (1993); Lastand Stevens (1994); Bonfil (1995, 1997); Menni, Hazin and Lessa (1995); Pollard, Smith, and Smith (1995);Villavicencio-Garayzar (1996); Santos, Porteiro and Barreiros (1997); Sheehan (1998); I.K. Fergusson, L.J.V. Compagno,K.J. Graham, F. Fakhoury, W. Noshie and W. Noshie (unpublished data); R. Bonfil (pers. comm.); D. Perrine (pers. comm.);C. Roessler (pers. comm.).Odontaspis noronhai (Maul, 1955) Fig. 56Carcharias noronhai Maul, 1955, Notul. Nat. Acad. Nat. Sci. Philadelphia, (279): 3, figs. 1-4. Holotype: Museu Municipal doFunchal, Madeira, MMF-2691, 1 710 mm female, from off Camara de Lobos, Madeira, between 600 and 1 000 m depth andmost likely at 800 to 1 000 m depth.Synonyms: None.Other Combinations: None.FAO Names: En - Bigeye sand tiger; Fr - Requin noronhai; Sp - Solrayo ojigrande.UPPER AND LOWER TEETH ON LEFT SIDEUNDERSIDE OF HEADFig. 56 Odontaspis noronhaiField Marks: A large, bulky shark with a long bulbously conical snout, eyes very large without nictitating eyelids, mouth longand extending behind eyes, teeth moderately large with prominent narrow cusps and a single pair of lateral cusplets, upperanterior teeth separated from lateral teeth by one or two rows of small intermediate teeth, anal fin and second dorsal finsmaller than first dorsal fin but broad-based, first dorsal fin on back and closer to pectoral fins than to pelvic fins, upperprecaudal pit present but lateral keels absent from caudal peduncle, caudal fin asymmetrical but with a strong ventral lobe,colour uniform blackish, dark chocolate brown or reddish brown, without spots, first dorsal fin with a light apical blotch.

Sharks of the World, Vol. 2 67Diagnostic Features: One row (occasionally two rows) of small intermediate teeth between upper anterior and lateral toothrows; a pair of rows or a single row of upper symphysial teeth present, but sometimes absent; usually 2 to 4 pairs of lowersymphysial tooth rows present; tooth rows 34 to 43/37 to 46 (71 to 83 total); root lobes moderately arched and broad;anterolateral teeth with a single pair of lateral cusplets. Pectoral fin rounded. First dorsal-fin apex broadly rounded in adult.Anal fin with height 2.4 to 4.5% of total length. Anal fin with straight posterior margin. Caudal fin with ventral caudal lobehardly developed. Colour: background colour of surface black, chocolate brown or dark reddish brown dorsally andventrally, without dark spots on sides of body; fins black, first dorsal fin often with a white blotch at tip.Distribution: Possibly circumglobal in all warm seas but as presently known sporadically distributed with very few records inthe Atlantic and Central Pacific. Western North Atlantic: USA (Texas, Gulf of Mexico). Central Atlantic off Mid-Atlantic Ridgejust north of the Equator. Western South Atlantic: Southern Brazil. Eastern North Atlantic: Madeira. ?Western Indian Ocean:possibly Seychelles. Pacific: South of Hawaiian Islands, USA; off Marshall Islands, possibly South China Sea.Habitat: Apparently an inhabitant of the continental and insular slopes near the bottom at 600 to 1 000 or more metres andwell off it near the surface at 100 m in water 640 m deep, and in the epipelagic and mesopelagic zone of the open ocean at 60to 450 m in water 4 500 to 5 300 m deep. Its uniform dark coloration, similar to many deepwater oceanic teleosts, suggests amesopelagic rather than epipelagic habitat, and it could be primarily an oceanic shark rather than a primarily slope-dwellingepibenthic species.Biology: This is a poorly known deepwater shark. One was caught at night near the Marshall Islands at 75 m with a longlineset over water between 4 500 and 5 300 m deep, suggesting that it may migrate vertically to the epipelagic zone at night anddescend in the daytime but this cannot be confirmed at present. It may also be migratory and seasonal in distribution, asBrazilian catches were made only during the springtime. Reproduction unknown, feeding habits little known; one specimenhad squid beaks and fish otoliths in its stomach. One caught near Hawaii writhed and snapped vigorously when captured.Size: Maximum at least 3.6 m; males possibly adolescent at 217 cm, three males adult at 326 to 342 cm long; a femaleimmature at 321 cm and another adult at about 326 cm.Interest to Fisheries and Human Impact: Incidental and apparently rare bycatch of pelagic longliners but little-utilized. Theholotype was taken on a vertical longline set by fishermen for black scabbardfish (Aphanopus carbo). Presumably takenoccasionally by deepwater fisheries with line and net gear, including pelagic gill nets, purse seines and deep-set longlines. Itmay live mostly below the depths normally fished by horizontal pelagic longlines and purse seines, and is possibly too largeto be a regular bottom or pelagic trawl catch. Conservation status unknown.Local Names: Bigeye sand tiger, Black sand tiger, Oceanic sand tiger.Remarks: Compagno (1981a) recognized this species on the type description (Maul, 1955) but suggested that it possiblywas only an extreme variant of O. ferox. However, descriptions of additional specimens convinced the writer that it is a validspecies, readily separable from O. ferox (Compagno, 1984; Sadowsky et al., 1984; Branstetter and McEachran, 1986;Humpreys, Moffitt and Seki, 1989). Some characters used by Compagno (1984), including the position of the first dorsal-finorigin, more vertical position of the first dorsal-fin margin, and position of the second dorsal-fin origin relative to the pelvic-finbases, apparently do not hold due to individual variation in O. noronhai and O. ferox.click for next page

click for previous page68 FAO Species Catalogue for Fishery Purposes No. 1A set of jaws possibly from the Seychelles (D. Ward, pers. comm.) has the dentitional characters of this species, and is thebasis for the Indian Ocean record of O. noronhai. Sadowsky et al. (1984) indicated that a possible alternative locality for thisjaw set was the South China Sea.Literature: Maul (1955); D’Aubrey (1964b); Sadowsky, Amorim and Arfelli (1984); Compagno (1984); Quero (1984);Branstetter and McEachran (1986); Humphreys, Moffitt and Seki (1989); Springer (1990); McEachran and Fechhelm(1998); D. Brogan (pers. comm.); D. Guitart-Manday (pers. comm.); V. Sadowsky (pers. comm.).2.2.2 Family MITSUKURINIDAEFamily: Family Mitsukurinidae Jordan, 1898, Proc. Calif. Acad. Sci. ser. 3 (Zool.), 1: 201.Type Genus: Mitsukurina Jordan, 1898.Number of Recognized Genera: 1.Synonyms: Family Scapanorhynchidae White, 1936: 4. Type genus: Scapanorhynchus Woodward, 1889, a Cretaceousfossil genus (see remarks below).FAO Names: En - Goblin sharks; Fr - Requins lutin; Sp - Tiburones duende.Diagnostic Features: Head as long as trunk or slightly shorter. Snout greatly elongated, blade-like and flattened. Eyessmall, length 1.0 to 2.4% of precaudal length. Gill openings short, length of first 4.6 to 5.9% of precaudal length, notextending onto dorsal surface of head; all gill openings anterior to pectoral-fin bases; no gill rakers on internal gill slits. Mouthlarge, parabolic, ventral on head; jaws strongly protrusable to about opposite snout tip but not greatly distensible laterally.Teeth large, anteriors and laterals very narrow and awl-like, in 35 to 53/31 to 62 (66 to 115 total) rows; three rows of largeanterior teeth on each side of upper and lower jaws, the uppers separated from the smaller upper lateral teeth by a gapwithout intermediate teeth; a pair of lower symphysial teeth present. Trunk compressed and moderately slender, very softand flabby. Caudal peduncle compressed and without keels or precaudal pits. Dermal denticles small and rough, with erectspike-like crowns with narrow cusps and ridges; cusps of lateral denticles pointing perpendicular to surface of skin. Pectoralfins short and broad, much shorter than head in adults; pectoral skeleton aplesodic with radials confined to fin bases. Pelvicfins large, larger than dorsal fins; fin skeleton aplesodic. Dorsal fins small, low, and rounded, or semi-angular, first andsecond dorsals equal-sized and smaller than the large, rounded anal fin; first dorsal skeleton aplesodic. Second dorsal andanal fins with broad, nonpivoting bases. Caudal fin not lunate, dorsal lobe long but half length of rest of shark or less, ventrallobe not developed. Neurocranium low, with a greatly elongated compressed rostrum, depressed internasal septum andwidespread nasal capsules, small orbits with the supraorbital crests reduced to isolated preorbital and postorbitalprocesses, tiny stapedial foramina, and with hyomandibular facets not extended outward. Vertebral centra strongly calcified,with well-developed double cones and radii but no annuli. Total vertebral count 122 to 125, precaudal count 53 to 56,diplospondylous caudal count 68 to 69. Intestinal valve of ring type with 19 turns. Size large, with adults 2.6 to 3.6 m long.Local Names: Goblin sharks, Chien wên sha k’o; Mitsukurizame-ka (Japan); Akuly domovye (Russia).Remarks: The living genus Mitsukurina has often been synonymized with the Upper Cretaceous fossil genusScapanorhynchus Woodward, 1889 (Woodward, 1899; Regan, 1906a; Goodrich, 1909; Engelhardt, 1913; Garman, 1913;White, 1936, 1937; Berg, 1940; Fowler, 1941; Romer, 1945, 1966; Bigelow and Schroeder, 1948; Berg and Svedovidov,1955; Matsubara, 1955; Garrick and Schultz, 1963; Norman, 1966; Bass, D’Aubrey and Kistnasamy, 1975a), but the two areapparently generically distinct (Signeux, 1949; Arambourg and Bertin, 1958; Glikman, 1967; Cappetta, l980, 1987;Compagno, 1973, 1984). The type species of Scapanorhynchus, S. lewisi (Davis, 1887) is known from whole-bodiedspecimens that show many differences from Mitsukurina owstoni (Cappetta, 1987). Thus Scapanorhynchus lewisi hasangular dorsal fins, a very long, low subangular anal fin, a more angular, much larger pectoral fin, a caudal fin with a discreteventral caudal lobe and a more angular terminal lobe, more normal denticles with flattened, expanded tricuspidate crowns,and somewhat different teeth with less expanded and flattened root lobes and often stouter cusps. Some palaeontologists(Glikman, 1967; Shelton P. Applegate (pers. comm.)) place Scapanorhynchus in a separate family (Scapanorhynchidae)from Mitsukurina, but others (Herman, 1977; Cappetta, 1980, 1987; Carroll, 1988) include them both in the Mitsukurinidae.Despite dentitional and other differences, the presence of a greatly expanded, paddlefish-like rostrum, anteriorly extendedsubethmoid fossa, elongated anal fin, and bent palatine processes on the palatoquadrates in both genera is stronglysuggestive of a relationship between them.Jordan (1898) proposed a new family, Mitsukurinidae, for the living goblin shark, which was recognized by severalsubsequent authors (Jordan, 1923; Lozano y Rey, 1928; Whitley, 1940; Glikman, 1967; Compagno, 1973, 1981b, 1984,1999; Carroll, 1988; Nelson, 1994; Shirai, 1996; Helfman, Collette and Facey, 1997; Eschmeyer, 1998). Following White(1936, 1937), many workers used the family Scapanorhynchidae for the goblin shark (Fowler, 1941, 1947, 1967a, b; Romer,1945; Bigelow and Schroeder, 1948; Matsubara, 1955; Budker and Whitehead, 1971; Lindberg, 1971; Rass and Lindberg,1971; Pinchuk, 1972; Bass, D’Aubrey and Kistnasamy, 1975a; Nelson, 1976; Gubanov, Kondyurin and Myagkov, 1986), butMitsukurinidae Jordan 1898 has priority (Compagno, 1984). Cappetta (1987) and Carroll (1988) also include the fossil

Sharks of the World, Vol. 2 69genus Anomotodon Arambourg, 1952 in the Mitsukurinidae, and include as a synonym of Mitsukurinidae the familyAnomotodontidae Herman, 1979. Several authors (Regan, 1906a; Engelhardt, 1913; Garman, 1913; Bertin, 1939a; Berg,1940; Fowler, 1941; Berg and Svedovidov, 1955; Arambourg and Bertin, 1958; Norman, 1966; Romer, 1966; Patterson,1967; Blot, 1969; Nelson, 1984; Eschmeyer, 1990), included Mitsukurina as a separate genus or as a synonym ofScapanorhynchus in the family Odontaspididae or Carchariidae, while Goodrich (1909) included it in the Lamnidae.Mitsukurina shows a curious mix of numerous primitive and derived characters that isolate it from other living lamnoids.Compagno (1990b), using morphological characters for a phyletic analysis of Lamniformes, and Long and Waggoner(1996), using dental morphology, suggested that Mitsukurinidae is the sister group of all other living Lamniformes. Recentmolecular phyletic studies (Naylor et al., 1997; Martin and Naylor, 1997) have not given unambiguous results on therelationship of Mitsukurinidae to other lamnoids, but one possibility is that Mitsukurinidae is sister to all other taxa.Morphological and molecular studies apparently support the retention of Mitsukurina in its own family.This family includes only a single living species, Mitsukurina owstoni, along with fossil species of Anomotodon,Mitsukurina and Scapanorhynchus. Morphological definition of the family is based on the living species.Mitsukurina Jordan, 1898Genus: Mitsukurina Jordan, 1898, Proc. Calif. Acad. Sci. ser. 3 (Zool.), 1: 199.Type Species: Mitsukurina owstoni Jordan, 1898, by monotypy.Number of Recognized Species: 1.Synonyms: None.Diagnostic Features: See family Mitsukurinidae above.Mitsukurina owstoni Jordan, 1898 Fig. 57Mitsukurina owstoni Jordan, 1898, Proc. Calif. Acad. Sci. ser. 3 (Zool.), 1: 200, pls. 11-12. Holotype: Zoological Museum,University of Tokyo, 107 cm immature male, near Yokohama, Japan, in deep water. Holotype lost, according to Eschmeyer(1998, Cat. Fish.: CD-ROM).Synonyms: Odontaspis nasutus de Braganza, 1904: 49, 104, pl. 1, figs. 1-1c. Type locality, Mare de Sezimbra, Portugal,603 m. Types unknown according to Eschmeyer (1998). Scapanorhynchus jordani Hussakof, 1909: 257, text-figs., pl. 44.Syntypes (2): American Museum of Natural History, AMNH-00004SW, jaws, model on display from 1 300 mm female;1 155 mm female, formerly in the Zoological Department at Columbia University. Type locality, Japan. Scapanorhynchusdofleini Engelhardt, 1912: 644. Holotype: Zoologischen Staatssammlung München, 2 100 mm female, Mayegawa, SagamiSea, Japan. Locality of holotype unknown according to Eschmeyer (1998: CD-ROM). Scapanorhynchus mitsukurii White,1937: 29 (error for Mitsukurina owstoni Jordan, 1898). Japan.Other Combinations: Scapanorhynchus owstoni (Jordan, 1898).FAO Names: En - Goblin shark; Fr - Requin lutin; Sp - Tiburón duende.UPPER AND LOWER TEETH ON RIGHT SIDE, LINGUAL VIEWFig. 57 Mitsukurina owstoniUNDERSIDE OF HEAD

70 FAO Species Catalogue for Fishery Purposes No. 1Field Marks: This unmistakable shark has a flat blade-like elongated snout, tiny eyes without nictitating eyelids, soft flabbybody, slender very long-cusped teeth in long highly protrusable jaws, two spineless dorsals and an anal fin, and a longcaudal fin without a ventral lobe. Colour: live and newly-captured individuals are pinkish white, but usually fade to brownishin alcohol.Diagnostic Features: See family Mitsukurinidae above.Distribution: Western Atlantic: Guiana, Surinam, French Guyana. Eastern Atlantic: France (Bay of Biscay), Madeira,Portugal, Senegal, Gulf of Guinea, South Africa (Western Cape). Western Indian Ocean: South Africa (Eastern Cape,KwaZulu-Natal), Mozambique (Mozambique seamount range). Western Pacific: Japan, Australia (South Australia, NewSouth Wales), New Zealand. Eastern Pacific: USA (southern California).Habitat: A poorly known, bottom-dwelling shark that inhabits the outer continental shelves and upper slopes and is found offseamounts, but rarely occurs at the surface or in shallow water close inshore. Most records are on or near the continentalslopes between 270 and 960 m deep but down to at least 1 300 m, sometimes in shallower shelf waters at 95 to 137 m.Seamount records suggest that the species is oceanic or semioceanic in addition to its known occurrences off continentalslopes.Biology: Very little is known of the biology of this bizarre shark, which is rare in most places where it is known apart fromJapan and possibly Portugal. The long flexible caudal fin, without a ventral lobe, the soft, flabby body, and small, soft pairedand unpaired fins, suggest that the goblin shark is a relatively inactive, slow swimming species with a density close toseawater. Its remarkable blade-like snout is superficially similar to those of the chondrostean paddlefishes (Polyodontidae),and like these fishes may use it as a forward-projecting prey detector. Its slender, pick-like anterior and lateral teeth suggestssmall, soft-bodied prey including fishes, shrimp and squid, and one specimen was found with fish remains in its stomach. Asin Carcharias taurus, the posterior teeth of the goblin shark are modified for crushing.The jaws of the goblin shark are highly specialized for rapid projection from the head as in some mesopelagic teleosts,propelled in part by a double set of elastic tensioning ligaments at the mandibular joints. The first set of ligaments are at thehinge joint between the ceratohyal head and Meckel’s cartilage on each side; and the second set extends across the head ofthe hyomandibula in a cavity between the ceratohyal and Meckel’s cartilage on each side. The ligaments are stretched whenthe jaws are retracted rearward into the mouth but are relaxed when the jaws are shot forward, and apparently function(along with the long preorbitalis muscles) like a catapult to project the jaws forward and snap up small animals.This shark is often illustrated and preserved with the jaws more or less protruded but a live goblin shark in captivity in theMarine Science Museum, Tokai University, Shimizu, Japan, held its jaws tightly retracted while swimming (Shiobara, 1990,Y. Shiobara, pers. comm. and photographs). Early catch records (Bean, 1905) suggested that mature females visited theeast coast of Honshu during the springtime only. Mode of reproduction unknown; a pregnant female has never beenreported.Size: Maximum total length at least 384 cm. Size at birth unknown, smallest recorded specimen 107 cm; mature males 264,320 and 384 cm, females reaching 373 cm, one mature at 335 cm. Weight 210 kg at 384 cm.

Sharks of the World, Vol. 2 71Interest to Fisheries and Human Impact: Interest to fisheries minimal, taken as untargeted bycatch of deepwater trawlfisheries and occasionally taken with deepwater longlines, deep-set gill nets, and possibly purse seines. Utilizeddried-salted for human consumption. Harmless to people. A spectacular aquarium exhibit, but seldom kept in captivity; onelived for a week in an aquarium at Tokai University, Shimizu, Japan. Conservation status unknown.Local Names: Nasuta (Portugal), Elphin or Elfin shark, Japanese goblin shark, Tenguzame or Tengu (goblin) shark,Mitsukurizame or Mitsukuri’s shark (Japan); Kabouterhaai (South Africa).Literature: Jordan (1898); Bean (1905); Hussakof (1909); Garman (1913); Fowler (1941); Bigelow and Schroeder (1948);Stead (1963); Bass, D’Aubrey and Kistnasamy (1975a); Uyeno, Nakamura and Mikami (1976); Piotrovsky and Prut’ko(1980); Uyeno, Matsuura and Fujii (1983); Cadenat and Blache (1981); Quero (1984); Compagno (1984); Davison and vanBerkel (1985); Stevens and Paxton (1985); Shcherbachev (1987); Compagno, Ebert and Smale (1989); Springer (1990);Shiobara (1990); Compagno, Ebert and Cowley (1991); Last and Stevens (1994); Shinohara and Matsuura (1997);P. Duarte (pers. comm.); D.A. Ebert (pers. comm.); Y. Shiobara (pers. comm.); J.D. Stevens (pers. comm.); J. Ugoretz (pers.comm.).2.2.3 Family PSEUDOCARCHARIIDAEFamily: Family Pseudocarchariidae Compagno, 1973, J. Linn. Soc.(Zool.), 53, suppl. 1: 28.Type Genus: Pseudocarcharias Cadenat, 1963.Number of Recognized Genera: 1.Synonyms: Family Pseudocarcharinidae Shirai, 1996: 34. Probably error for Pseudocarchariidae.FAO Names: En - Crocodile sharks; Fr - Requins crocodile; Sp - Tiburones cocodrilo.Diagnostic Features: Head much shorter than trunk. Snout moderately long, pointed and bulbously conical, not greatlyelongated or flattened and blade-like. Eyes very large, length 3.6 to 4.9% of precaudal length. Gill openings moderately long,length of first 5.4 to 8.2% of precaudal length, extending onto dorsal surface of head; all gill openings in front of pectoral-finbases; no gill rakers on internal gill slits. Mouth large, parabolic, ventral on head; jaws strongly protrusable to almostopposite snout tip but not greatly distensible laterally. Teeth large, the anteriors narrow and awl-like, the laterals morecompressed and blade-like, with 26 to 29/21 to 26 (45 to 52 total) rows; two rows of enlarged anterior teeth on each side ofupper jaw, the uppers separated from the smaller upper lateral teeth by a row of small intermediate teeth; three rows of loweranteriors on each side, the first two rows enlarged but the third about as large as laterals; symphysials absent. Trunkcylindrical and slender. Caudal peduncle slightly depressed and with low lateral keels and upper and lower crescenticprecaudal pits present. Dermal denticles small and smooth, with flat crowns, small ridges and cusps, and with cuspsdirected posteriorly on lateral denticles. Pectoral fins small, short and broad, much shorter than head in adults; pectoralskeleton aplesodic with radials confined to fin bases. Pelvic fins large, somewhat smaller than pectoral and first dorsal fins;fin skeleton aplesodic. First dorsal fin small, low, and angular; fin skeleton aplesodic. Second dorsal fin smaller than first butlarger than anal fin; second dorsal fin with a broad nonpivoting base but anal fin pivotable. Caudal fin not lunate, dorsal lobemoderately long but less than half as long as rest of shark, ventral lobe short but strong. Neurocranium moderately high, witha short to moderately elongated rostrum, depressed internasal septum and narrowly separated nasal capsules, large orbitswith the supraorbital crests strong, small stapedial fenestrae, and with hyomandibular facets not extended outward.Vertebral centra strongly calcified, with well-developed double cones and radii but no annuli. Total vertebral count 146 to158, precaudal count 80 to 88, diplospondylous caudal count 60 to 71. Intestinal valve of ring type with 24 to 27 turns. Sizesmall with adults 0.74 to 1.10 m.Local Names: Crocodile sharks, Tubarões crocodilos (Mozambique).Remarks: The single living genus and species in this family, Pseudocarcharias kamoharai, was formerly placed in thefamily Odontaspididae and genus Odontaspis or Carcharias (Matsubara, 1936; Teng, 1959; D’Aubrey, 1964a, b; Bass,D’Aubrey and Kistnasamy, 1975a). Anatomical work by the writer has shown that this species is very distinct and rates aseparate family Pseudocarchariidae. Characters of the family are presented and discussed in Compagno (1973, 1977,1982, 1984, 1990b, 1999). Pseudocarchariidae has been recognized by a number of subsequent writers (Last and Stevens,1994; Nelson, 1994; Shirai, 1996; Helfman, Collette and Facey, 1997). Eschmeyer (1990) reduced its rank to a subfamilyPseudocarchariinae of the family Odontaspididae, but subsequently recognized it as a full family (Eschmeyer, 1998).Pseudocarcharias was proposed by Cadenat (1963) as a subgenus of Carcharias Rafinesque, 1810 but was synonymizedwith Carcharias by D’Aubrey (1964a, b) and Odontaspis Agassiz, 1838 by Bass, D’Aubrey and Kistnasamy (1975a). It isstrongly divergent from either Odontaspis or Carcharias as delimited here and has been recognized by many writers (Abeet al., 1969; Compagno, 1973, 1977, 1981a,b, 1982, 1984, 1990b, 1999; Krefft, 1980; Fujita, 1981; Cadenat and Blache,1981; Nakaya, 1984; Springer, 1990; Cigala-Fulgosi, 1992; Last and Stevens, 1994). The synonymy of the species followsD’Aubrey (1964a, b), Bass, D’Aubrey and Kistnasamy (1975d) and Compagno (1973, 1984).

72 FAO Species Catalogue for Fishery Purposes No. 1Compagno (1990b) suggested, on morphological characters, that Pseudocarchariidae might be the sister group ofMegachasmidae, Alopiidae, Cetorhinidae, and Lamnidae within Lamniformes. Recent molecular phyletic studies (Naylor etal., 1997; Martin and Naylor, 1997) have not given unambiguous results on the relationship of Pseudocarchariidae to otherlamnoids, but suggest relationships with Odontaspididae and Megachasmidae. Long and Waggoner (1996), using dentalmorphology for a phyletic analysis of Lamniformes, suggested that Pseudocarchariidae is the immediate sister group ofAlopiidae plus Lamnidae within the Lamniformes.Pseudocarcharias Cadenat, 1963Genus: Subgenus Pseudocarcharias Cadenat, 1963 (Genus Carcharias Rafinesque, 1810), Bull. Inst. Francaise AfriqueNoire, ser. A, 25(2): 526 (proposed as a subgenus of Carcharias Rafinesque, 1810, but used throughout in generic form).Type Species: Pseudocarcharias pelagicus Cadenat, 1963, by original designation, a junior synonym of Carchariaskamoharai Matsubara, 1936.Number or Recognized Species: 1.Synonyms: None.Diagnostic Features: See family Pseudocarchariidae above.Pseudocarcharias kamoharai (Matsubara, 1936) Fig. 58Carcharias kamoharai Matsubara, 1936, Zool. Mag. Tokyo, 48(7): 380. Holotype: Imperial Fisheries Institute, Japan, KyotoUniversity, Department of Fisheries, Faculty of Agriculture, Japan (housed at Maizuru, Japan) FAKU, Fish Spec. 1823,735 mm male, Koti Fish Market, Koti, Japan, apparently lost according to Eschmeyer (1998, Cat. Fish.: CD-ROM).Synonyms: Carcharias yangi Teng, 1959: 1, fig. 1. Holotype, Taiwan Fisheries Research Institute, TFRI 2895, 1 000 mmTL adult male, Su-ao fish market, from off Su-ao, Taiwan (Province of China). Type status confirmed by Eschmeyer (1998:CD-ROM). Pseudocarcharias pelagicus Cadenat, 1963: 529, figs. 1-5. Holotype: Museum National d’Histoire Naturelle,Paris, MNHN 1963-1, 975 mm adult male, off the Guinea coast, West Africa. Type status confirmed by Eschmeyer (1998:CD-ROM).Other Combinations: Odontaspis kamoharai (Matsubara, 1936).FAO Names: En - Crocodile shark; Fr - Requin crocodile; Sp - Tiburón cocodrilo.UPPER AND LOWER TEETH ON RIGHT SIDE, LINGUAL VIEWFig. 58 Pseudocarcharias kamoharaiUNDERSIDE OF HEAD

Sharks of the World, Vol. 2 73Field Marks: A small, very distinctive oceanic shark, with huge eyes lacking nictitating eyelids, long gill slits, slender,spindle-shaped body, long-cusped prominent teeth in a long angular mouth with highly protrusable jaws, small pectoral fins,two small spineless dorsal fins and an anal fin, weak keels and precaudal pits on the caudal peduncle, an asymmetricalcaudal fin with a long ventral lobe. Colour: grey or grey-brown dorsal surface, lighter ventral surface, and light-edged fins.Diagnostic Features: See family Pseudocarchariidae above.Distribution: Oceanic and circumtropical. Western Atlantic: Off Brazil. Eastern Atlantic: Southeast of Cape Verde Islands,between them and Guinea-Bissau, Guinea, Angola, and South Africa (Western Cape, vicinity of Cape Town and CapePeninsula). Western Indian Ocean: Mozambique Channel between southern Madagascar, southern Mozambique andKwaZulu-Natal, South Africa, possibly south within Agulhas Current to off Eastern Cape. ? Eastern Indian Ocean: Bay ofBengal (possibly erroneous). Western North Pacific: Off Japan, Taiwan Island and Korean Peninsula; area betweenMarshall, Howland and Baker, Palmyra, Johnston and Hawaiian Islands. Western South Pacific: Australia (northeasternQueensland), west of New Zealand (North Island), Coral Sea, Indonesia (south of Sumatra near Sunda Straits and off Java).Central Pacific: Marquesas Islands, Hawaiian Islands, open ocean between Marquesas and Hawaiian Islands, open oceanbetween Hawaiian Islands and Baja California, around Line Islands, open water between Line Islands and southern Peru.Eastern Pacific: Mexico (off west coast of Baja California), Costa Rica, Panama and northern Peru.Habitat: A rare to locally abundant oceanic, epipelagic and possibly mesopelagic shark, usually found offshore and far fromland but sometimes occurring inshore and near the bottom, at depths from the surface to at least 590 m. Its bicolorate,countershaded colour pattern, lack of an expanded iris and prominent green or yellow retinal reflection, and frequentoccurrence in pelagic longline catches suggests that it primarily inhabits the epipelagic zone. There are several records ofstrandings in the Cape Town area, South Africa, possibly due to upwelling of cold water that may stun these sharks but in atleast one instance as a discarded catch of an offshore longliner.Biology: The long body cavity, large liver, and small fins of this shark (microceanic habitus, Compagno, 1990a) give it asuperficial resemblance to Isistius, Squaliolus, Euprotomicrus, Scymnodalatias, and other oceanic squaloids, as well asOdontaspis noronhai, and like these sharks its extremely large and oily liver is probably important in maintaining neutralbuoyancy. Its habits are little known, but its firm body musculature, tough skin, small precaudal fins, and large caudal finsuggests that it is a relatively active species, which is also suggested by its behaviour when captured. Off Cape Point, SouthAfrica, one jumped out of the water after a bait and was caught. It snaps strongly and vigorously when captured (S. Kato,pers. comm.) and can bite very hard. The large but nonreflective eyes of the crocodile shark suggest nocturnal activity in theepipelagic zone, and possibly a diel pattern of movement toward the surface at night and away from it in the day.The crocodile shark is ovoviviparous and a uterine cannibal, with the young having yolk sacs at 3 to 4 cm long butreabsorbing them and subsisting on eggs and possibly other young beyond this size. Number of young in a litter four, two peruterus; egg cases formed in the oviducts have 2 to 9 fertilized eggs, but apparently only two of these survive, possiblythrough elimination of extra rivals. An interesting question is why two young survive in each uterus in this shark and someother lamnoids, while in Carcharias taurus only one foetus per uterus is normally produced.Feeding habits of this shark are sketchily known. Its long, flexed teeth, strong and long jaws, and its vigorous activity whencaptured adapt it to moderately large, active oceanic prey. Of seven specimens examined by the writer for stomach

74 FAO Species Catalogue for Fishery Purposes No. 1contents, the stomachs of four were empty and three others had a number of small bristlemouths (gonostomatids), possiblylanternfish (myctophids), unidentified fish scales, small shrimp, and squid beaks, including onychoteutids (Moroteuthisrobsoni), mastigoteuthids (Mastigoteuthis), pholidoteuthids (Pholidoteuthis ?boschmai), and cranchiids(Megalocranchia?) in their stomachs (M. Roeleveld and M. Lipinski, pers. comm., on identification of squid beaks). Thejaws of the crocodile shark can be protruded for a considerable distance from its head.Size: This is the smallest living lamnoid, with maximum size at least 110 cm; size at birth about 41 cm; males adult at 74 to110 cm; adolescent females examined 96 to 110 cm and adults recorded at 89 to 110 cm and presumably greater.Interest to Fisheries and Human Impact: This shark is primarily caught as a discarded bycatch of pelagic longlinefisheries for scombroids, but details are sketchy. Abe et al. (1969) noted that the species is often caught on tuna longlines,but discarded because of its small size and meat that is apparently unsuitable for the Japanese market. The liver of thisspecies is very large and very high in squalene, and hence is of potential value. It also has been caught on squid jigs andoccasionally washes up on beaches in the Cape Town area of South Africa. It may also be a discarded bycatch of pelagicsquid fisheries as well as of pelagic net fisheries for scombroid fishes.The crocodile shark is small and has never bitten people in the water, but should be treated with respect because of itsstrong jaws. It apparently has not been kept by large public aquaria but its small size and poorly-known behaviour suggests itmay be more amenable to captivity than larger pelagic lamnoids and might be an interesting animal to observe in captivity.Conservation status is uncertain but of concern because of its epipelagic habitat and because it is an apparentlywidespread, discarded, and largely unrecorded bycatch of the burgeoning pelagic longline fisheries. It is too small to be ofmuch value for fins, and is little-utilized for flesh, but has a large mouth and strong teeth and is readily caught on longlinehooks fished near the surface. It does not appear to be abundant anywhere with the known exception of the MozambiqueChannel in the western Indian Ocean during the 1960s, and catch records are very limited and largely confined to a smallnumber of specimens (less than 50) deposited in museums. It was assessed as Limited Risk (Near Threatened) for the RedList of the IUCN Shark Specialist Group (L.J.V. Compagno and J.A. Musick, pers. comm.).Local Names: Kamohara’s sand shark, Mizu-wani, Mizuwani, Water crocodile, Water alligator (Japan); Japanese raggedtooth shark, Grootoog-skeurtandhaai, Bigeye ragged-tooth (South Africa).Literature: Matsubara (1936);Teng (1959); Lindberg and Legeza (1959); Cadenat (1963); Chen (1963); D’Aubrey (1964a,b); Merrett (1965); Abe et al. (1969); Abe (1973); Bass, D’Aubrey and Kistnasamy (1975a); Compagno (1973, 1981, 1982,1984, 1990a, b); Krefft (1980); Fujita (1981); Cadenat and Blache (1981); Nakaya (1984); Sadowsky, Amorim and Arfelli(1987); Kashkin (1989); Fischer et al. (1990); Springer (1990); Cigala-Fulgosi (1992); Last and Stevens (1994); Long andSeigel (1996); L.J.V. Compagno and J.A. Musick (pers. comm.); S. Kato, (pers. comm.).2.2.4 Family MEGACHASMIDAEFamily: Family Megachasmidae Taylor, Compagno and Struhsaker, 1983, Proc. California Acad. Sci., ser. 4, 43(8): 87, 89.Type Genus: Megachasma Taylor, Compagno and Struhsaker, 1983.Number of Recognized Genera: 1.Synonyms: None.FAO Names: En - Megamouth sharks; Fr - Requins grande gueule; Sp - Tiburones bocudos.Diagnostic Features: Head very elongated and about length of trunk. Snout extremely short, flattened and broadlyrounded, not elongated and blade-like. Eyes moderately large, length 1.6 to 1.8% of precaudal length. Gill openingsmoderately long, length of first 6.4 to 8.6% of precaudal length, not extending onto dorsal surface of head; last two gillopenings over pectoral-fin bases; unique gill rakers of finger-like dermal papillae with cartilage cores fringing internal gillslits. Mouth very large and arcuate, terminal on head; jaws greatly protrusable anteriorly beyond snout tip but not greatlydistensible laterally. Teeth small, continuously varying and more or less awl-shaped, in 55 to 115/75 to 121 (130 to 236 total)rows, no differentiation between anterior, intermediate, lateral, or symphysial teeth. Trunk cylindrical and somewhatcompressed, stout, and relatively flabby. Caudal peduncle compressed and without lateral keels but with smallfossate-shaped upper precaudal pit only. Dermal denticles very small and smooth, with flat crowns, small ridges and cuspsand with cusps directed posteriorly on lateral denticles. Pectoral fins large, narrow and elongated, much shorter than head inadults; pectoral skeleton plesodic with radials extending far into fin webs. Pelvic fins moderate-sized, smaller than pectoraland first dorsal fins; fin skeleton probably aplesodic, not extending into fin web. First dorsal fin moderately large, semierectand angular; fin skeleton aplesodic. Second dorsal fin less than half size of first but moderately large. Anal fin smaller thansecond dorsal fin and with its base slightly behind second dorsal-fin base, bases of both fins not pivotable. Caudal fin not

Sharks of the World, Vol. 2 75lunate, dorsal lobe long but less than half as long as rest of shark, ventral lobe short but strong. Neurocranium depressed,with short wide rostrum, greatly depressed internasal septum and widespread nasal capsules, small orbits with strongsupraorbital crests, small stapedial fenestrae, and with hyomandibular facets not extended outward. Vertebral centra weaklycalcified, with rudimentary radii and double cones and no annuli. Total vertebral count 151, precaudal count 64,diplospondylous caudal count 82 to 87. Intestinal valve of ring type with 23 or 24 turns. Size very large with adults 4.5 to5.5 m long.Remarks: Taylor, Compagno and Struhsaker (1983) proposed a new monotypic family of lamnoid sharks, Megachasmidae,for the new genus and species of megamouth shark (Megachasma pelagios), and suggested that it might be either the sistergroup of all other lamnoids or could be grouped with more derived lamnoids having plesodic pectoral fins. The familyMegachasmidae has received recognition from various authors, including Compagno (1984, 1990b, 1999), Gubanov,Kondyurin and Myagkov (1986), Eschmeyer (1990, 1998), Nelson (1994), Last and Stevens (1994), Shirai (1996), andHelfman, Collette and Facey (1997). Maisey (1985) adopted the second suggestion of the megamouth shark being relatedto derived plesodic lamnoids, but maintained that the megamouth shark was the sister group of the basking shark andconfamilial with it in the family Cetorhinidae on derived characters of its jaw suspension, cranial morphology, dentition andfilter-feeding. Robins et al. (1991a) followed Maisey’s classification. Compagno (1990b) noted that Maisey’s derivedcharacters for grouping the basking and megamouth sharks did not hold, and listed numerous characters separating thesesharks. He suggested on morphological grounds that the Megachasmidae was distinct from Cetorhinidae, that Cetorhinidaewas the immediate sister group of Lamnidae, that Megachasmidae was a valid taxon, that filter-feeding was independentlyderived in the basking and megamouth sharks, and that the Megachasmidae, on the basis of plesodic pectoral fins, was thesister group of all other plesodic lamnoids (Alopiidae, Cetorhinidae and Lamnidae). Two molecular studies confirmed theseparation of the Megachasmidae and Cetorhinidae and refuted the common evolution of filter-feeding in these sharks, butdisagreed on the relationships of Megachasmidae within the Lamniformes. Martin and Naylor (1997), using cytochrome bmitochondrial genes, suggested that Cetorhinus was sister to the Lamnidae but could not resolve the position ofMegachasma robustly except for its arising from the basal root of lamnoids; their work suggested no common grouping ofplesodic lamnoids but a possible relationship of Megachasma to Pseudocarcharias plus Odontaspis. Morrissey, Dunn andMulé (1997) using the 12S messenger RNA mitochondrial gene suggested that Megachasma was the sister of all otherlamnoids that they examined. Long and Waggoner (1996), using dental morphology for a phyletic analysis of Lamniformes,suggested that the basking and megamouth sharks were immediate sister groups within the Lamniformes, and formed asister taxon to Odontaspididae, Pseudocarchariidae, Alopiidae, and Lamnidae. As noted in Compagno (1990b), similaritiesin tooth morphology between these sharks could be the result of parallelism and may not reflect a common immediate originwithin the Lamniformes. Particularly problematical is the absence of disjunct monognathic heterodonty in Megachasma,while Cetorhinus retains disjunct upper anteriors and a gap between laterals and anteriors as in all other lamnoids. Althoughthe phylogeny of Lamniformes remains unsettled, morphological and molecular studies support the retention of theMegachasmidae as distinct from the Cetorhinidae or any other lamnoid family.Megachasma Taylor, Compagno and Struhsaker, 1983Genus: Megachasma Taylor, Compagno and Struhsaker, 1983, Proc. California Acad. Sci., ser. 4, 43(8): 87, 96.Type Species: Megachasma pelagios Taylor, Compagno and Struhsaker, 1983, by original designation.Number of Recognized Species: 1.Synonyms: None.Diagnostic Features: See family Megachasmidae above.Megachasma pelagios Taylor, Compagno and Struhsaker, 1983 Fig. 59Megachasma pelagios Taylor, Compagno and Struhsaker, 1983, Proc. California Acad. Sci., ser. 4, 43(8): 87, 96, figs. 1-15.Holotype: Bernice P. Bishop Museum, BPBM-22730, 4 460 mm adult male, off Oahu, Hawaiian Islands, 21° 51’ N,157° 46’ W, about 42 km northeast of Kahuku Point, at 165 m depth in water about 4 600 m deep.Synonyms: None.Other Combinations: None.

76 FAO Species Catalogue for Fishery Purposes No. 1FAO Names: En - Megamouth shark; Fr - Requin grande gueule; Sp - Tiburón bocudo.Field Marks: One of three species of gigantic filter-feeding non-batoid sharks, unmistakable with its extremely short butbroadly rounded snout, very large and long head, huge terminal mouth that extends behind the eyes, numerous smallhooked teeth, moderately long gill slits, internal gill slits lined with dense rows of papillose gill rakers, eyes without nictitatingeyelids, two dorsal fins and an anal fin, caudal peduncle without keels, caudal fin asymmetrical not lunate, but with a shortand strong ventral lobe, and no light spots. Colour: upper surface of body grey or grey-black above, without light spots,underside white, mouth dusky blackish with dark spotting on lower jaw, and dorsal surfaces of pectoral and pelvic finsblackish with conspicuous light margins.Diagnostic Features: See family Megachasmidae above.Fig. 59 Megachasma pelagiosDistribution: Spottily known from less than 20 specimens but probably circumtropical and wide-ranging. Western Atlantic:Brazil. Eastern Atlantic: Senegal. Southeastern Indian Ocean: Australia (Western Australia). Western North Pacific: Japan,Philippines (Macabalan Bay, Cagayan de Oro), Indonesia (Sulawezi, Nain Island, Bunaken Archipelago). Central Pacific:Hawaiian Islands (Oahu). Eastern Pacific: USA (southern California near San Clemente and off Catalina Island and SanDiego).Habitat: A coastal and oceanic, epipelagic and neritic species, found in water as shallow as 5minashallow bay and inwater 40 m deep on the continental shelf, with at least one washed ashore alive on a sandy beach; also offshore in theepipelagic zone at 8 to 166 m depth in water 348 to 4 600 m deep. The coloration and catch records of the megamouth sharkare suggestive of epipelagic rather than deepwater habitat, as is the composition of its liver oil (Itabashi, Yamaguchi andNakaya, 1997).

Sharks of the World, Vol. 2 77Biology: This is a seldom reported, possibly rare or uncommon shark, with most examples from off Japan and southernCalifornia. As the megamouth shark has greatly reduced teeth, very numerous gill-raker papillae on its internal gill openings,and stomach packed with very small prey, it can be properly considered a specialized filter feeder like the basking shark(Cetorhinus maximus), whale shark (Rhincodon typus), and the devil rays (Mobulidae). However, the flabby body, soft fins,asymmetrical caudal fin without keels, and weak calcification of the megamouth shark suggested that it is much less activethan the whale and basking sharks (Compagno, 1984, 1990b) and the mobulids. Observations and tracking of a livespecimen off southern California confirmed its relative sluggishness (Nelson et al., 1997).The only known prey of the megamouth shark are epipelagic and mesopelagic euphausiid shrimp, copepods, and jellyfish.The first megamouth shark from Hawaii had been feeding on an euphausiid shrimp, Thysanopoda pectinata, that averages3.1 cm in length. The shrimp has a diel migration pattern with a range of 300 to 1 100 m depth during the day; at night it iscommonest at 150 to 500 m with a maximum range of 75 to 525 m. When captured during the night the shark was apparentlyat the upper depths where these shrimp are commonest. It may have been feeding on them when it fouled itself by its mouthand teeth on a pair of parachutes being used as sea anchors by a US Navy research vessel. The second megamouth sharkfrom California had euphausiids, copepods and deepwater jellyfish (Atolla vanhoeffeni) in its stomach. Two female sharksfrom Japan had macerated euphausiids in their stomachs (probably Euphausia nana). The feeding structures of this sharkmay allow it to feed on other pelagic invertebrates and even small midwater fishes, but so far the limited stomach contentsavailable suggests that this shark primarily targets euphausiid shrimp as prey.There has been considerable speculation on the feeding habits of this shark. Taylor, Compagno and Struhsaker (1983)compared the basking, whale, and megamouth sharks, and suggested that the megamouth shark slowly swam throughschools of euphausiid shrimp with jaws widely open, occasionally closing its mouth and contracting its pharynx toconcentrate prey before swallowing it. These authors noted the presence of a bright silvery mouth lining, which theysuggested was possibly bioluminescent but could not prove it because of inadequate histological evidence. The secondspecimen had tissue in its lower jaw that may be luminescent (Lavenberg and Seigel, 1985) and could along with thereflective upper jaw tissue serve as a ‘light trap’ for luring prey to the proximity of its mouth (Diamond, 1985). Compagno(1990b) noted that the protrusability of the jaws in the megamouth shark may allow it to use its mouth as a bellows to suck inprey, and that open-mouthed cruising through prey concentrations was optimal for the basking shark but not the megamouthshark.Taylor, Compagno and Struhsaker (1983) suggested that the soft flabby body and fins, low-flow filter apparatus, and smallgill openings of the megamouth shark indicated that it was less active than the whale and basking sharks. Oikawa andKanda (1997) considered gill histology and filament area in the megamouth shark compared to the shortfin mako and otherspecies, and suggested that the megamouth shark was less active. An adult male megamouth shark of about 4.9 m lengthwas captured in a pelagic gill net off southern California near the surface in water between 300 and 400 m deep. This wasstill alive when discovered in the net and was towed to Dana Point Harbor where it was tethered by a rope to the fishing boatthat caught it (Nelson et al., 1997). The shark was extensively photographed by divers and was still alive after being towedand then being tethered for over a day. Observations suggested that the shark could breathe readily by gill pumping and wasnot dependent on ram-ventilation and constant swimming unlike more active lamnoid sharks. The shark was then towed outto sea, tagged with an acoustic telemetric tag, and released in water about 20 m deep. The shark immediately descended,outswam the accompanying divers, and headed for deep water beyond the continental shelf. It was tracked over a two-dayperiod, which revealed a pattern of vertical, crepuscular migration in the epipelagic zone. The tagged shark ascended atsunset to a depth of 12 to 25 m at night, then descended at sunrise to a depth of 120 to 166 m during the day with greatestdepth achieved at midday but stayed well above the bottom at 700 to 850 m. It stayed on a straight southward course at anestimated speed between 1.5 and 2.1 km/hr during day and night. It was thought that the shark was responding to light levelsin choosing its depth range, and as light increased it correspondingly sought an optimal level by diving at dawn andconversely responded to decreasing light levels by ascending to near (but not at) the surface at dusk. It has been suggestedthat the megamouth shark may also follow vertical migrations of euphausiid prey during diel cycles. The telemetric tracksuggested that the shark was indeed somewhat less active than makos or basking sharks, but that it could sustain a slowrate of swimming for extended periods.Mode of reproduction is probably aplacental viviparous with uterine cannibalism or cannibal vivipary suspected in the form ofoophagy, but no pregnant adult female has been reported to date. A late immature or early adolescent female had twoovaries with many tiny oocytes under 3 mm in diameter, while an adult female had numerous larger oocytes 5 to 10 mmwide. This is similar to the ovaries of several other lamnoids that are oophages. An adult female had numerous bite marksmostly on her flanks and precaudal tail but also on her first dorsal and anal fin and head. These wounds corresponded to theteeth of another megamouth shark (or sharks), and were interpreted as courtship scars inflicted by a male because of thenarrow spacing of individual cuts comparable to male rather than female teeth (Yano et al., 1999).The megamouth shark is the only known selachian victim of the semiparasitic cookiecutter shark, Isistius brasiliensis, andmay be especially vulnerable to Isistius attacks because of its soft skin, epipelagic habitat in warm seas where Isistius isalso found, sluggishness, and relatively slow swimming speed. Three megamouth specimens had ‘crater wounds’ indicativeof cookiecutter attacks. A megamouth shark was seen at the surface off Nain Island, Bunaken Archipelago, North Sulawesiwhile being possibly harassed or played with by three sperm whales, which left the shark after the observers approachedthem. The shark was apparently minimally injured by the whales and was photographed at the surface before disappearing.

78 FAO Species Catalogue for Fishery Purposes No. 1Size: Maximum total length at least 549 cm. Size at birth unknown. A juvenile free-living male from off Brazil was 190 cmlong while a possibly smaller specimen from off Senegal was estimated at about 180 cm long. Adult males were 446 to about549 cm; a late immature or early adolescent female was 471 cm, another female of uncertain maturity was about 5 m, and anadult female was 544 cm.Interest to Fisheries and Human Impact: Interest to fisheries minimal at present. Taken as a rare incidental bycatch inpelagic gill nets, purse seines, pelagic longlines, and fixed shore nets, and so far has been mostly utilized by museums andoceanaria, which prize the few specimens landed as display objects. As with certain other large, rare animals it attractsmuch attention from the general public and shark fans. A few specimens were released alive from fishing gear, while a recentspecimen from Philippines was cut up and utilized by fishermen, but details of its utilization were not recorded. Conservationstatus uncertain, but of concern because of the apparent epipelagic and neritic habitat and possible rarity of this shark, whichputs it at risk as unrecorded bycatch of oceanic and offshore littoral fisheries.Local Names: Megamouth sharks, Big mouth sharks.Remarks: This giant pelagic filter-feeding shark is perhaps the most spectacular discovery of a new shark in the twentiethcentury. A recent symposium volume deals primarily with a detailed examination of a single specimen of megamouth sharkfrom Japan, but includes papers on all aspects of megamouth biology (Yano et al., 1997).Literature: Taylor, Compagno and Struhsaker (1983); Compagno (1984, 1990b); Lavenberg and Seigel (1985); Diamond(1985); Maisey (1985); Nakaya (1989); Berra and Hutchins (1990, 1991); Miya, Hirosawa and Mochizuki (1992); Last andStevens (1994); Clark and Castro (1995); Carey and Clark (1995); Seret (1995); Nelson et al. (1997); Castro et al. (1997);Berra (1997); Yano et al. (1997); Morrissey, Dunn and Mulé (1997); Martin and Naylor (1997); Nakaya et al. (1997);Yabumoto et al. (1997); Oikawa and Kanda (1997); Tanaka and Yano (1997); Itabashi, Yamaguchi and Nakaya (1997);Yamaguchi and Nakaya (1997); Yano et al. (1999); E.T. Elizaga (pers. comm.); E. Gomes, A. Amorim and B. Hueter (pers.comm.); H. Mollet (pers. comm.); J. Morrissey (pers. comm.); P. Pecchioni (pers. comm.); D. Petersen (pers. comm.).2.2.5 Family ALOPIIDAEFamily: Subfamily Alopiadini Bonaparte, 1838 (Family Squalidae), Nuov. Ann. Sci. Nat., Bologna, ser. 1, 2: 209. Emendedto Family Alopiidae Bonaparte, 1838 by Jordan and Gilbert (1883, Bull. U.S. Nat. Mus., 16: 26).Type Genus: Alopias Rafinesque, 1810.Number of Recognized Genera: 1.Synonyms: Family Alopeciae Müller and Henle, 1839: 74. Type genus: Alopecias Müller and Henle, 1837. FamilyVulpeculidae Garman, 1913: 12, 30. Type genus: Vulpecula Garman, 1913.FAO Names: En - Thresher sharks; Fr - Renards; Sp - Zorros.Field Marks: Long curving asymmetrical caudal fin with dorsal lobe nearly or quite as long as rest of shark, short ventralcaudal lobe, long narrow pectoral fins, large pelvic fins, large first dorsal fins, tiny second dorsal and anal fins, large to hugeeyes.Diagnostic Features: Head much shorter than trunk. Snout moderately long, pointed and conical, niether greatlyelongated, nor flattened and blade-like. Eyes moderately large to very large with length 1.8 to 4.3% of precaudal length. Gillopenings short, with width of first 3.1 to 5.2% of precaudal length, not extending onto dorsal surface of head; last two pairs ofgill openings over pectoral-fin bases; no gill rakers on internal gill slits. Mouth small and arcuate, ventral on head, jaws notstrongly protrusable. Teeth small to moderately large, compressed and blade-like, in 22 to 52/20 to 50 (42 to 102 total) rows.Two rows of small to moderately large anterior teeth on each side of upper jaw, the anteriors hardly larger than adjacentlaterals and separated from them by a row of small intermediate teeth or a gap; three rows of small lower anterior teeth oneach side, these slightly larger than lower laterals and with the third row not reduced in size; small symphysial teeth presentor absent in upper and lower jaws. Trunk cylindrical and moderately stout, firm and not flabby. Caudal peduncle slightlycompressed, with upper and lower crescentic precaudal pits but no lateral keels. Dermal denticles very small and smooth,with flat crowns, small ridges and cusps and with cusps directed posteriorly on lateral denticles. Pectoral fins very long andnarrow, longer than head in adults; pectoral skeletons plesodic with radials extending far into fin webs. Pelvic fins very large,nearly or quite as large as first dorsal fin; fin skeleton plesodic. First dorsal fin large, high, erect and angular; fin skeletonsemiplesodic, extending partially into fin web. Second dorsal and anal fins minute, low and with pivoting bases, anal-fin basebehind second dorsal-fin base. Caudal fin not lunate, dorsal lobe greatly elongated, about as long as rest of shark, ventrallobe short but strong. Neurocranium high and compressed, with short rostrum, compressed internasal septum and closely

Sharks of the World, Vol. 2 79adjacent nasal capsules, huge orbits with well-developed supraorbital crests, small stapedial fenestrae, and withhyomandibular facets not extended outward. Vertebral centra strongly calcified, with well-developed double cones and radiibut without prominent annuli. Total vertebral count 282 to 477, precaudal count 100 to 125, diplospondylous caudal count180 to 356. Intestinal valve of ring type with 33 to 45 turns. Size moderately large to very large with adults 2.7 to 5.5+ m long.Distribution: Threshers occur worldwide in tropical, subtropical, and warm and cold-temperate waters.Habitat: Threshers are large, active, strong-swimming sharks, ranging in habitat from coastal to epipelagic and deepwaterepibenthic.Biology: Thresher sharks are large, active sharks that are apparently specialized for feeding on small to moderately largeschooling fishes and squids. Threshers swim in circles around a school of prey, narrowing the radius and bunching theschool with their long, strap-like caudal fins. The caudal fin is also used as a whip to stun and kill prey, and threshers arecommonly tail-hooked on longlines after striking the bait with the caudal tip. The three known species of this family broadlyoverlap in habitat and range, but differences in their structure, feeding habits and spatial distribution suggest that theyreduce interspecific competition by partitioning their habitat and available prey to some extent. Alopias superciliosus, withits huge eyes, relatively large teeth, broad caudal fin, and preference for deeper water (including the outer shelves and upperslopes near the bottom), takes somewhat larger pelagic fishes (including small billfishes and lancetfishes) as well as bottomfishes. A. vulpinus, with smaller eyes and teeth, a narrower caudal fin, and preference for the surface and coastal as well asoceanic waters, takes small pelagic fishes (including clupeids, needlefishes and mackerels) and squids, but also bonitosand bluefishes. The oceanic and marginally coastal A. pelagicus is less well known biologically, but its much smaller teethand very slender caudal fin suggest that it may take smaller prey than A. vulpinus or A. superciliosus. Evidence on vascularmorphology and from telemetry data suggests that threshers are endothermic (Alexander, 1998), but their behaviour(including activity patterns) is poorly known despite their abundance and wide range.Interest to Fisheries and Human Impact: Thresher sharks form an important component of oceanic and offshore coastalshark fisheries, particularly because of their high-quality meat which is utilized fresh, frozen, smoked and dried-salted. Theirfins are used for shark-fin soup, livers for vitamin extraction, and hides for leather. Sizeable oceanic fisheries for threshersharks, utilizing floating longlines, have operated in the northwestern Indian Ocean, the central Pacific, the western Pacific,and the western North Atlantic. Threshers are undoubtedly caught as bycatch of longline fisheries targeting scombroidfishes. A pelagic gill net fishery for threshers has operated off the Pacific coast of the USA over the last few decades butdeclined markedly due to overfishing. Threshers are also captured offshore and near shore with line gear (including rod-andreel)and fixed bottom gill nets. Sports anglers seek threshers as game fishes, because of their strong fighting qualities andtendency to jump when hooked. Little is available on fisheries statistics for thresher catches worldwide. At present, only NewZealand and the USA report commercial catch statistics of thresher sharks to FAO (A. vulpinus) and these amount to onlyabout 100 t or less annually during the last decade. World catches are undoubtedly much larger. Threshers are sometimesseen by divers, but do not, to the writer’s knowledge, provide ecotouristic viewing at inshore or offshore dive sites, nor havethey been regularly kept in aquaria. The conservation status of threshers is poorly known, but is of concern because of theirhigh value in fisheries, very low fecundity, and their occurrence in areas and habitats subjected to high-intensity oceanicfisheries.Local Names: Fox sharks, Threshers, Thresher sharks, Thrashers (English); Renards de mer, Chienhai chang, Chienhaichang wei sha k’o; Dlinnokhvostye akuly (Russia); Onagazame-ka (Japan); Zorros (Mozambique).Remarks: Following Müller and Henle (1839), most authors have recognized the threshers (genus Alopias) as a separatefamily, Alopiidae or equivalents, which is followed here. However, some authors placed the threshers in the family Lamnidaeor Isuridae (Günther, 1870; Hasse, 1879; Woodward, 1889; Regan, 1906a; Goodrich, 1909; Engelhardt, 1913; Berg, 1940;Berg and Svetovidov, 1955; Arambourg and Bertin, 1958; and Norman, 1966). Shirai (1996) recently included the baskingsharks (Cetorhinidae) and mackerel sharks (Lamnidae) as subfamilies of the Alopiidae.The arrangement of genera and species within this family follows Bass, D’Aubrey and Kistnasamy (1975a), Gruber andCompagno (1981), and Compagno (1984) in recognizing a single living genus and three living species, Alopias pelagicus,A. superciliosus and A. vulpinus. Compagno (1990b), on external and cranial morphology, suggested that threshers weremonophyletic, with A. vulpinus the plesiomorphic sister group of A. pelagicus and A. superciliosus. Eitner (1995) didelectrophoretic analysis of allozymes from muscle samples collected from eastern Pacific threshers. Eitner suggested asimilar phyletic relationship as indicated by Compagno (1990b) for the three known species, but indicated that a fourth andpossibly undescribed species, initially identified as A. superciliosus, occurred off Baja California, Mexico. Unfortunately thefourth species was not represented by whole-bodied material, parts or even photographs or morphometrics (only musclesamples), so that determination of its status awaits collection and description of additional material from the eastern Pacific.The three known species of threshers are strongly differentiated by external and skeletal morphology (Compagno, 1990b),so that detailed morphological examination of the possible fourth species is highly desirable.Literature: Garman (1913); Fowler (1941, 1967a); Bigelow and Schroeder (1948); Garrick and Schultz (1963); Lindberg(1971); Shiino (1972, 1976); Compagno (1973, 1984, 1988, 1990a, b, 1999); Bass, D’Aubrey and Kistnasamy (1975a);Gruber and Compagno (1981); Last and Stevens (1994); Eitner (1995); Alexander (1998).

80 FAO Species Catalogue for Fishery Purposes No. 1Alopias Rafinesque, 1810Genus: Alopias Rafinesque, 1810, Caratt. gen. sp. anim. piant. Sicilia, Palermo, pt. 1: 13.Type Species: Alopias macrourus Rafinesque, 1810, by monotypy, a junior synonym of Squalus vulpinus Bonnaterre,1788, Tabl. Encyclop. Method. Trois Reg. Nat., Ichthyol., Paris: 9.Number of Recognized Species: 3.Synonyms: Genus Vulpecula Jarocki, 1822: 454. Probably based on Vulpecula marina Valmont, 1768 (work notconsistently binomial), equivalent to Squalus vulpinus Bonnaterre, 1788. Genus Alopecias Müller and Henle, 1837a: 114.Type species: “Carcharias vulpes Cuv[ier]” by original designation, equals Squalus (Carcharias) vulpes Cuvier, 1816.Genus Alopius Swainson, 1838: 91 (unjustified emendation of Alopias Rafinesque, 1810). Genus Vulpecula Garman,1913: 3, 30. Type species: Vulpecula marina Garman, 1913, by monotypy: “Valmont, 1768, gives a description of V.marina of earlier authors. His species is Squalus vulpinus Bonn., 1788, the Alopias macrourus Raf., 1810, A. vulpesBonap. 1841. The genus and the species are adopted from Valmont” (Garman, 1913: 3). Revival of Vulpecula Valmont(1768: 740). Valmont’s names were rejected as being inconsistently binomial by the International Commission on ZoologicalNomenclature (1925, Opinion 89: 27-33). Genus Alopes Vladykov and McKenzie, 1935: 46 (erroneous spelling for Alopias).Key to Species:1a. Head nearly flat between eyes, with a deephorizontal groove on nape on each side abovegills; eyes very large, with orbits expanded ontodorsal surface of head (Fig. 60); teeth large, in 22to 27 rows in upper jaw; first dorsal-fin base closerto pelvic bases than to pectoral bases . . . . .. . . . . . . . . . . . . . . . . . . . Alopias superciliosus1b. Head strongly arched between eyes, without ahorizontal groove or with an inconspicuous oneon nape on each side; eyes smaller, with orbitsnot expanded onto dorsal surface of head(Fig. 61); teeth smaller, 32 to 52 rows in upper jaw;first dorsal-fin base about equidistant betweenpectoral and pelvic-fin bases or closer topectoral-fin bases . . . . . . . . . . . . . . . . . . . 2deep grooveLATERAL VIEW OF HEADFig. 60 Alopias superciliosusLATERAL VIEW OF HEADFig. 61 Alopias pelagicus2a. Head narrow, snout more elongated, foreheadnearly straight; labial furrows absent; pectoral finsnearly straight and broad-tipped; sides abovepectoral bases dark, without an extension of thewhite abdominal area (Fig. 62) . . . . . Alopias pelagicus2b. Head broad, snout shorter, forehead stronglyarched; labial furrows present; pectoral finsbroad tippedfalcate and narrow-tipped; sides abovepectoral-fin bases marked with a white patchextending forward from the abdominal area(Fig. 63) . . . . . . . . . . . . . . . . . Alopias vulpinusno white patchFig. 62 Alopias pelagicuswhite patchnarrow tippedFig. 63 Alopias vulpinusclick for next page

click for previous pageSharks of the World, Vol. 2 81Alopias pelagicus Nakamura, 1935 Fig. 64Alopias pelagicus Nakamura, 1935, Mem. Fac. Sci. Agric. Taihoku Imp. Univ. ,14(1): 2, 3, pl. 1, fig. 1. Syntypes: Three largeindividuals mentioned and measured, these 270, 285, and 330 cm TL, and a large female specimen illustrated but ofuncertain size; also foetus 96.5 cm (Nakamura, ibid., 5, pl. 3), probably referable to Alopias vulpinus; all from Suô fishmarket, Taiwan (Province of China). Whereabouts of syntypes uncertain according to Eschmeyer (1998, Cat. Fish.:CD-ROM).Synonyms: None.Other Combinations: Alopias vulpinus (not Bonnaterre, 1788).FAO Names: En - Pelagic thresher; Fr - Renard pélagique; Sp - Zorro pelágico.UPPER AND LOWER TOOTHUNDERSIDE OF HEADFig. 64 Alopias pelagicusField Marks: Long upper caudal lobe nearly as long as rest of shark, relatively small eyes, very narrow head with archeddorsal profile, straight broad-tipped ‘oceanic’ pectoral fins, first dorsal fin somewhat closer to pectoral-fin bases thanpelvic-fin bases, very slender caudal-fin tip, body colour deep blue or grey above, white below, white colour of abdomen notextending over pectoral-fin bases.Diagnostic Features: Head narrow in dorsal and ventral views, with a convex, arched dorsolateral profile. Snoutmoderately long, conical. Eyes moderately large in adults but very large in newborn and foetuses, not expanded onto dorsalsurface of head and without a vertical, binocular field of view; interorbital space broadly convex. Labial furrows absent. Teethvery small, in 41 to 45/37 to 38 rows (total for both jaws 75 to 86 rows); posterior tooth rows 5 to 11; symphysial andintermediate tooth rows usually present. Weak nuchal grooves present above branchial region. Pectoral fins of‘macroceanic’ type with straight and very broad tips. Claspers moderately slender and not whip-like. First dorsal-fin midbaseabout equidistant between pectoral and pelvic-fin bases or closer to pectoral-fin bases. Caudal tip very slender with verynarrow terminal lobe. Ribs of monospondylous precaudal vertebrae fused ventrally to form a canal extending nearly to theocciput. Total vertebral count 453 to 477. Intestinal valve count 37 to 40. Colour: body deep blue to grey on upper surfacewith sides silvery and underside white, white colour of abdomen not extending over pectoral-fin bases; no white dot on upperpectoral-fin tips.Distribution: Oceanic and wide-ranging in the Indo-Pacific. Indian Ocean: South Africa (KwaZulu-Natal), Red Sea, Gulf ofAden, Arabian Sea (off Somalia, between Oman and India, and off Pakistan), Australia (northwest Western Australia),Western North Pacific: China, Taiwan (Province of China), Japan (southeastern Honshu). Western South Pacific: NewCaledonia, eastern Micronesia, Tahiti. Central Pacific: Hawaiian Islands, equatorial waters north of Howland and Baker,Phoenix and Palmyra Islands. Eastern Pacific: USA (California) and Mexico (Baja California, Gulf of California), equatorialwaters northwest of French Polynesia, and off Galapagos Islands.

82 FAO Species Catalogue for Fishery Purposes No. 1Habitat: Primarily an oceanic, epipelagic, circumtropical species, but sometimes caught near shore on beaches with anarrow continental shelf, ranging in depth from the surface to at least 152 m. Sometimes seen by divers near coral reefs,near dropoffs and in large lagoons, and on sea mounts.Biology: A little-known, active, strong-swimming species, probably migratory but with movements little-known. In theeastern North Pacific there is a possible population centre off central Baja California, which tends to shift northward (alongwith other oceanic sharks) during strong El Niño events. Behaviour and sociobiology is poorly known. Michael (1993) hasseen this species repeatedly leap (breach) out of the water.Ovoviviparous, with uterine cannibalism as in other species of Alopias. Embryos subsist on their yolk-sacs up to about12 cm, after which they become oophagous, feeding on unfertilized eggs. No evidence of adelphophagy (embryo-eating)was reported by Liu et al. (1999), who examined 233 embryos from 167 pregnant females. Litter size is two, with one foetusper uterus and with sex ratio 1:1. Gestation period uncertain because females give birth all year long without a definite birthseason. Liu et al. suggest that the gestation period may be less than a year as with A. vulpinus, but because most adultfemales were pregnant throughout the year there may be an annual cycle with no resting period between pregnancies.Pupping may also occur in winter in the Gulf of Aden (R. Bonfil, pers. comm.). This species presumably feeds on small fishesand squid but no details are known.Vertebral growth rings are laid annually in vertebral centra; females mature at about 8 or 9 years old and males at about 6 to9 years old, with up to 16 growth rings for females and 14 for males for a minimal age of 14 to 16 years old and a maximumage estimated from von Bertalanffy growth curves as 20 years for males and 29 years for females. Assuming birth of twoyoung every year a female might produce about 40 young during her lifetime. This species has unusually large young, withthe largest known foetus 43% of the length of the largest adult female. The large size of the young may help to reducepostnatal predation (presumably by other large sharks), but the relatively small size of the adults combined with the lowfecundity imposed by large foetal size may in turn require annual breeding.Size: Maximum total length at least 365 cm. Size at birth uncertain but presumably between about 130 and 160 cm andpossibly up to 190 cm. The largest term foetus examined by Liu et al. (1999) off Taiwan (Province of China) was 158 cm andtheir smallest specimen was 190 cm long and a year old; a freeliving specimen from the western Indian Ocean that wasexamined by the author was 137 cm long. A term or near-term foetus 96.5 cm long attributed to this species by Nakamura(1935) is probably A. vulpinus. Off Taiwan (Province of China) males were immature at about 174 to 283 cm, adolescent atabout 239 to 305 cm, and adult at 259 to 323 cm; onset of maturity was at about 267 cm, with 50% mature at 267 to 276 cm.Females from Taiwan (Province of China) were immature at 176 to 294 cm, adolescent at 253 to 321 cm, and adult at 265 toabout 365 cm; onset of maturity was at about 273 cm, with 50% mature at 282 to 292 cm. Elsewhere males were adolescentat 192 to 318 cm and adult at 276 cm, while females were immature or adolescent at 277 to 233 cm, adult at 264 to 330 cm,while pregnant females were 264 to about 300 cm. This is apparently a smaller species than A. superciliosus or A.vulpinus.Length-weight equations are given by Liu et al. (1999) for Taiwanese specimens:Females: W(kg) = 4.61 x 10 -5 TL(cm) 2.494 (n = 230)Males: W(kg) = 3.98 x 10 -5 TL(cm) 2.52 (n = 230)

Sharks of the World, Vol. 2 83Interest to Fisheries and Human Impact: This species was formerly exploited by the longline fishery in the northwesternIndian Ocean (primarily by the former USSR), but it is also fished in the Central Pacific. It is an important catch off Taiwan(Province of China) with about 222 t landed yearly. It enters the commercial thresher catch off California and is sometimescaught in considerable numbers off the Pacific coast of Mexico and Gulf of California. Also caught by shark fishermen in theRed Sea and Gulf of Aden (R. Bonfil, pers. comm.). Utilized for its meat (for human consumption), liver oil for vitamin-Aextraction, hides for leather, and fins for shark-fin soup. Apparently seldom caught by anglers, but listed as a record fishalong with other threshers by the International Game Fish Association. It is rarely caught by anti-shark nets offKwaZulu-Natal, South Africa. Harmless to people, seldom encountered by divers and not kept in aquaria as far as is known.Divers have viewed and photographed this shark on coral reefs and seamounts in the Gulf of California and the Red Sea,Indonesia and Micronesia, but it is not a regular subject of ecotouristic diving. According to Michael (1993) it is shy anddifficult to approach underwater. The conservation status of this shark is uncertain, but Liu et al. (1999) considered itextremely vulnerable to overexploitation and in need of close monitoring because of its very low fecundity and relatively highage at maturation.Local Names: Pelagic thresher, Thresher shark, Whiptail shark, Fox shark; Onagazame, Nitara (Japan); Zorro pelagico(Mozambique).Remarks: Nakamura (1935) described A. pelagicus from three large specimens 270 to 330 cm TL , for which he presentedmeasurements. He also gave an illustration (pl. 1, fig. 2) of a large female specimen, one-twentieth life size. It is uncertain ifthe specimen illustrated is one of the three large individuals described, as scaling up the drawing (204 mm long) gives a sizeof 408 cm. He also included a separate description (p. 5) and illustration (pl. 3) of a foetus 96.5 cm long as A. pelagicus. All ofthese specimens are apparently syntypes of A. pelagicus. Nakamura did not designate types for A. pelagicus and did notindicate if one of the three large specimens measured or the large female illustrated was the mother of the illustrated foetusor if the latter was separately obtained.The large A. pelagicus specimen illustrated by Nakamura is apparently conspecific with A. pelagicus of Bass, D’Aubrey andKistnasamy (1975a), Compagno (1984), and Last and Stevens (1994), as shown by its fin shapes, fin positions, colourpattern (including lacking a white patch over its pectoral base), oblique teeth, and possibly by lacking labial furrows.However, the illustrated foetus is apparently A. vulpinus and is recognizable by its small eyes, broad head with a stronglyconvex dorsal profile, short snout, presence of labial furrows, and falcate pectoral fins. Compagno (1984) and Eschmeyer(1998) were unable to provide information on whether or not the syntypes of this species still exist or for that matter if theywere even preserved and deposited in a research collection. Nakamura only mentioned that the specimens were drawn byhim from life. It may be necessary to designate a neotype for A. pelagicus based on Taiwan (Province of China) material.The name A. pelagicus is used here in the sense of Bass, D’Aubrey and Kistnasamy (1975a), Compagno (1984), and Lastand Stevens (1994), who served to revise the concept of this species by restricting it to the species represented byNakamura’s illustrated adult.Alopias pelagicus has commonly been mistaken for A. vulpinus. For example, Gohar and Mazhar (1964, Red Sea), Kato,Springer and Wagner (1967, eastern Pacific), Fourmanoir and Laboute (1976, New Caledonia), Johnson (1978, Tahiti), andFaughnan (1980, Hawaiian Islands) all published illustrations of this species under the name A. vulpinus. This species isprobably more wide-ranging than present records show, although it has not been found in the Atlantic Ocean norMediterranean Sea and may be absent there.Literature: Nakamura (1935); Gohar and Mazhar (1964); Kato, Springer and Wagner (1967); Fourmanoir and Laboute(1976); Johnson (1978); Faughnan (1980); Bass, D’Aubrey and Kistnasamy (1975a); Otake and Mizue (1981); Compagno(1984, 1990a, b); Hanan, Holts and Coan (1993); Michael (1993); Last and Stevens (1994); Eitner (1995);Villavicencio-Garaysar, Estrada-Agüero and Downton-Hoffman (1997); Liu et al. (1999); R. Bonfil (pers. comm.); S.P.Applegate (pers. comm.); S. Kato (pers. comm.); J. Crow (pers. comm.).Alopias superciliosus (Lowe, 1840) Fig. 65Alopecias superciliosus Lowe, 1840, Proc. Zool. Soc. London, 1840(8): 39 (1841?). Also Lowe, 1849, Trans. Zool. Soc.London, 3(1): 18 (sometimes dated 1839). Holotype unknown (Eschmeyer, 1998, Cat. Fish.: CD-ROM), type localityMadeira, eastern Atlantic.Synonyms: Alopias profundus Nakamura, 1935: 2, pl. 1, fig. 1, pl. 2. Syntypes: Three large specimens, 332, 352 and366 cm TL, a large female illustrated and of uncertain size (Nakamura, 1935, pl. 1, fig. 1); also a 72 cm foetus, presumablythe same as illustrated (Nakamura, 1935., pl. 2); all specimens from Suô fish market, Taiwan (Province of China).Whereabouts of syntypes unknown according to Eschmeyer (1998: CD-ROM), possibly never accessioned in a researchcollection.Other Combinations: None.

84 FAO Species Catalogue for Fishery Purposes No. 1FAO Names: En - Bigeye thresher; Fr - Renard à gros yeux; Sp - Zorro ojón.Fig. 65 Alopias superciliosusDORSAL VIEW OF HEADField Marks: Long dorsal caudal lobe nearly as long as rest of shark, notched or helmeted contour of head, huge eyesextending onto dorsal surface of head, falcate but rather broad-tipped pectoral fins.Diagnostic Features: Head broad in dorsal and ventral view, with a notched dorsolateral profile. Snout moderately long,bulbous. Eyes greatly enlarged in young and adults, expanded onto dorsal surface of head and with a vertical, binocular fieldof view; interorbital space nearly flat. Labial furrows absent. Teeth large and in 22 to 27/20 to 24 rows (total for both jaws 42to 51 rows); posterior tooth rows 1 to 3; no symphysial or intermediate teeth. Strong nuchal grooves present above branchialregion. Pectoral fins falcate with curved and moderately broad tips. Claspers moderately slender and not whip-like. Firstdorsal midbase closer to pelvic-fin bases than to pectoral-fin bases. Caudal tip broad with wide terminal lobe. Ribs ofmonospondylous precaudal vertebrae fused ventrally to form a canal extending nearly to the occiput. Total vertebral count219 to 319. Intestinal valve count 43 to 45. Colour: body purplish grey or grey-brown on upper surface and sides withunderside grey to white, light colour of abdomen not extending over pectoral-fin bases; no white dot on upper pectoral-fintips.Distribution: Oceanic and coastal, virtually circumglobal in tropical and temperate seas. Western Atlantic (including Gulf ofMexico): USA (Atlantic coast from New York to Florida, Gulf of Mexico off Florida, Mississippi and Texas), Mexico (Veracruzto Yucatan), Bahamas, Cuba, Venezuela, central and southern Brazil. Eastern Atlantic: Portugal, Spain, Madeira, nearAzores, Morocco, Canary Islands, Senegal, Guinea to Sierra Leone, Angola, South Africa (Western Cape): also westernand central Mediterranean Sea. Indian Ocean: South Africa (Eastern Cape and KwaZulu-Natal), Madagascar, Arabian Sea(Somalia), Gulf of Aden, Maldives, Sri Lanka. Western Pacific: Southern Japan (including Okinawa), Taiwan (Province ofChina), Viet Nam, between northern Mariana Islands and Wake Island, Northwestern Submarine Rise, New Caledonia,Australia (northwestern coast), New Zealand. Central Pacific: Area between Wake, Marshall, Howland and Baker, Palmyra,Johnston, and Hawaiian Islands; north and south of Hawaiian Islands, off east of Line Islands, and between Marquesas andGalapagos Islands. Eastern Pacific: USA (California), Mexico (Gulf of California) and west of Galapagos Islands (Ecuador);possibly off Peru and northern Chile.

Sharks of the World, Vol. 2 85Habitat: Found in coastal waters over the continental shelves, sometimes close inshore in shallow waters, and on the highseas in the epipelagic zone far from land; also caught near the bottom in deep water on the continental slopes. Ranges fromthe surface and in the intertidal to at least 500 m deep, but mostly below 100 m depth.Biology: An epipelagic, neritic, and epibenthic shark, apparently strong-swimming but little-known behaviourally.Ovoviviparous, with uterine cannibalism (oophagy), number of young usually two per litter, but sometimes with 3 or 4. Sexratio of foetuses 1:1. Larger females apparently do not have larger term foetuses than smaller females. Birth may occurthroughout the year although in the eastern Atlantic more females may give birth in autumn and winter than other times ofyear. The gestation period may be 12 months long but remains uncertain because of lack of birthing seasonality. In theeastern Atlantic a nursery area occurs off the Straits of Gibraltar, but similar areas no doubt occur elsewhere in the vastrange of this species. Bigeye threshers have been aged with annular growth rings in vertebral centra, assuming one ring peryear (Liu, Chiang and Chen, 1998). Males mature at about 9 or 10 years old and females at about 12 or 13 years old.Maximum number of growth bands, and maximum known age, about 19 for males and 20 for females.The bigeye thresher feeds on pelagic fishes, including lancetfishes (Alepisauridae), herring (Clupeidae), mackerel(Scombridae), and small billfishes (Istiophoridae) and bottom fishes including hake (Merluccidae); also squids(Ommastrephidae). Apparently this species stuns its prey with its long caudal fin, as individuals are often tail-hooked onlonglines. The arrangement of the eyes, with keyhole-shaped orbits extending onto the dorsal surface of the head, suggestthat this species has a dorsal, vertical binocular field of vision (unlike other threshers) which may be related to fixating onprey and striking them with its tail from below. Michael (1993) observed sea lampreys attached near the cloaca of a bigeyethresher.Size: Maximum total length about 461 cm. Size at birth 100 to 140 cm, with full term foetuses at 105 or 106 cm andfree-swimming individuals down to 155 cm. Males immature up to 316 cm, maturing at about 279 to 300 cm; adult males assmall as 276 cm and reaching about 410 cm with an estimated maximum of 421 cm. Females immature up to 350 cm, andmaturing at about 294 to 355 cm; adult females as short as 341 cm and reaching at least 458 cm. Length-weight equationsfor this species are given by Kohler, Casey and Turner (1995) for fork length:W (kg) = 9.1069 x 10 -6 x FL (cm) 3.0802( n = 55; both sexes)where FL (cm) = 0.5598 x TL (cm) + 17.666 (n = 56)Liu, Chiang, and Chen (1998) give equations for total length:Females: W (kg) = 1.02 x 10 -5 TL (cm) 2.78 (n = 175)Males: W (kg) = 3.73 x 10 -5 TL (cm) 2.57 (n = 65)Interest to Fisheries and Human Impact: Caught or formerly caught in the oceanic longline fisheries operated by theformer USSR, Japan, Taiwan (Province of China), Spain, the USA, Brazil, Uruguay, Mexico, and probably other countries.Especially important areas for these fisheries are the northwestern Indian Ocean, western and Central Pacific, easternNorth Pacific, and North Atlantic. The bigeye thresher was formerly a very important component of the Cuban longlinefishery, and more recently has been taken in considerable numbers by longliners off the northeastern USA and by gill netvessels off southern California (USA) and the eastern Atlantic (by Spanish vessels), and by longliners off Taiwan (Provinceof China; where about 220 t per year are landed). This species is also taken as incidental bycatch in fixed bottom and pelagicgill nets, in trawls, and as a rare catch of anti-shark nets off KwaZulu-Natal, South Africa. It has been caught by anglers withsportsfishing gear (rod-and-reel) in the USA, South Africa, and New Zealand, in some instances by anglers targetingswordfish at night and using luminous lures. It is listed as a record fish along with other threshers by the International GameFish Association. Its meat is utilized fresh, smoked and dried-salted for human consumption, its liver oil is processed forvitamin A, its skin for leather, and fins for shark-fin soup. Apparently harmless to people, and not known to have beenencountered by divers underwater.Local Names: Big-eyed thresher, Zorro (Cuba); Zorro ojón (Mexico); Tubarâo raposo, Bigeye thresher (Azores); Deepseabigeye thresher, Bigeye, Bigeyed thresher shark, Big-eyed thresher, Hachiware (Japan); Grootoog-sambokhaai (SouthAfrica); Zorro olho grande (Mozambique).Remarks: This account follows Bass, D’Aubrey and Kistnasamy (1975a), Gruber and Compagno (1981), and Compagno(1984) in synonymizing Alopias profundus with this species. See Gruber and Compagno (1981) for a detailed discussion ofthe synonymy of A. profundus and for a general review of the biology of A. superciliosus.Literature: Nakamura (1935); Springer (1943); Bigelow and Schroeder (1948); Cadenat (1956c); Fitch and Craig (1964);Bass, D’Aubrey and Kistnasamy (1975a); Stillwell and Casey (1976); Blasco and Chapuli (1981); Gruber and Compagno(1981); Cadenat and Blache (1981); Branstetter and McEachran (1983); Gilmore (1983, 1993); Fulgosi (1983); Compagno(1984, 1990b, 1994); Compagno and Smale (1986); Ivanov (1986); Moreno and Morón (1992a); Hanan, Holts and Coan(1993); Michael (1993); Last and Stevens (1994); Eitner (1995); Kohler, Casey and Turner (1995); Bonfil (1997); Santos,Porteiro and Barreiros (1997); Chen, Liu and Chang (1997); Castillo-Geniz et al. (1998); Liu, Chiang and Chen (1998).

86 FAO Species Catalogue for Fishery Purposes No. 1Alopias vulpinus (Bonnaterre, 1788) Fig. 66Squalus vulpinus Bonnaterre, 1788, Tabl. Encyclop. Method. Trois Reg. Nat., Ichthyol., Paris: 9. Types unknown accordingto Eschmeyer (1998, Cat. Fish.: CD-ROM), type locality, Mediterranean Sea.Synonyms: Squalus vulpes Gmelin, 1788: 1496. Types unknown according to Eschmeyer (1998: CD-ROM). Alopiasmacrourus Rafinesque, 1810: 12. Type locality, Sicily. No types. Galeus vulpecula Rafinesque, 1810: 13, equivalent to“Squalus vulpecula di Linnao”, which does not exist. Type locality, Sicily. No types. Squalus alopecias Gronow, 1854: 7. Notypes known, according to Eschmeyer (1998: CD-ROM). Alopecias barrae Perez Canto, 1886: 5. Type locality, Chile.Holotype possibly lost. Alopecias chilensis Philippi, 1902: 310. Type locality, Chile. No types known, according toEschmeyer (1998: CD-ROM). Alopecias longimana Philippi, 1902: 308. Type locality, Chile. Whereabouts of type unknown,according to Eschmeyer (1998: CD-ROM). Vulpecula marina Garman, 1913: 30, pl. 7, fig. 1. Holotype: Museum ofComparative Zoology, Harvard, MCZ 1166-S (Eschmeyer, 1998:CD-ROM), juvenile 1 321/1 346 mm long (52 or 53 in) or546 mm PCL, from Massachusetts Bay. Revival of Vulpecula marina Valmont, 1768: 740, rather than a description of a newspecies. Valmont’s names were rejected as being not consistently binomial by the International Commission on ZoologicalNomenclature (1925, Opinion 89: 27-33). Alopias caudatus Phillipps, 1932: 226. Based on specimen in MelbourneMuseum figured by McCoy (1885: pl. 88) from Victoria, Australia. Holotype possibly NMV 58434 or 58437 according toEschmeyer (1998: CD-ROM). Alopias greyi Whitley, 1937: 5. Holotype? ca. 305 cm, type locality, Bermagui, New SouthWales; jaws only, whereabouts unknown according to Eschmeyer (1998: CD-ROM).Other Combinations: Alopias vulpes (Gmelin, 1788), Alopecias vulpes (Gmelin, 1788), Carcharias vulpes (Gmelin,1788).FAO Names: En - Thresher shark; Fr - Renard; Sp - Zorro.Fig. 66 Alopias vulpinusField Marks: Long curving dorsal caudal lobe about as long as rest of shark, relatively small eyes, falcate pointed pectoralfins, white colour of abdomen extending over pectoral-fin bases.Diagnostic Features: Head broad in dorsal and ventral view, with a strongly convex dorsolateral profile. Snout relativelyshort, conical and pointed. Eyes moderately large at all sizes, not expanded onto dorsal surface of head and without avertical, binocular field of view; interorbital space broadly convex. Labial furrows present. Teeth smaller with 32 to 52/25 to50 rows (total for both jaws 58 to 102 rows); posterior tooth rows 2 to 10; symphysial and intermediate tooth rows usuallypresent. No nuchal grooves present above branchial region. Pectoral fins falcate and with curved and narrow tips. Claspersextremely slender and whip-like. First dorsal-fin midbase closer to pectoral-fin bases than to pelvic fin bases. Caudal tipmoderately slender with moderately broad terminal lobe. Ribs of monospondylous precaudal vertebrae lateral and not fusedventrally as a canal. Total vertebral count 339 to 364. Intestinal valve count 33 to 34. Colour: body blue-grey to dark grey orblackish above with sides silvery or coppery and underside white, white colour of abdomen extending dorsally and anteriorlyover pectoral-fin bases as a conspicuous patch; white dot often present on upper pectoral-fin tips.Distribution: Oceanic and coastal, virtually circumglobal in tropical to cold-temperate seas but commonest in temperatewaters. Western Atlantic (including Gulf of Mexico): Canada (Newfoundland, Nova Scotia, New Brunswick, and Quebec,north to the Bay of Chaleur, Gulf of St. Lawrence), USA (entire Atlantic Coast but rare south of New England; Gulf Coast offFlorida, Mississippi and Texas), Cuba, Mexico (Veracruz to Campeche), Venezuela, Brazil to Argentina. Eastern Atlantic:Norway and British Isles to Mediterranean and Black Seas, Morocco, Madeira, the Azores, Ghana, Côte d’Ivoire, Angola,Namibia, and South Africa (Western Cape and probably Northern Cape). Indo-West Pacific: South Africa (Eastern Cape andKwaZulu-Natal), Tanzania, Somalia, Maldives, Chagos Archipelago, Gulf of Aden, possibly Oman, Pakistan, India, SriLanka, Sumatra, Japan, Republic of Korea, China, Taiwan (Province of China), Australia (Queensland, New South Wales,Victoria, Tasmania, South Australia), New Zealand, New Caledonia. Central Pacific: Society Islands, Fanning Islands,Hawaiian Islands. Eastern Pacific: Canada (British Columbia), the USA (Washington, Oregon and California) and Mexico(Baja California), south to Panama and Chile.

Sharks of the World, Vol. 2 87Habitat: Coastal over the continental and insular shelves and epipelagic far from land in temperate to tropical waters, mostabundant near land; young often close inshore and in shallow bays. Depth range from the surface and the intertidal to atleast 366 m, often near the surface.Biology: An active, strong-swimming shark, sometimes leaping out of the water. Thresher sharks in the northwestern IndianOcean and off the west coast of North America show spatial and depth segregation by sex. Off the west coast of NorthAmerica (and probably elsewhere) the species is seasonally migratory, and moves northward from Baja California intoCalifornia waters during the spring, with adult males tending to travel farther northward than females and reaching the coastof British Columbia. Juveniles are mostly found in shallow warm-temperate inshore waters, particularly off southernCalifornia where an important nursery area occurs. Juveniles may be less cold-tolerant than adults, and seldom range northof central California. Both adults and juveniles congregate in inshore waters of southern California, primarily during springand summer. Behaviour is otherwise poorly known, and little is known of sociobiology and behaviour patterns.Transoceanic migrations have not been demonstrated, and there may be separate populations with slightly differentfecundity and size at maturity in the eastern Pacific and western Indian Ocean, and possibly elsewhere, but this remains tobe determined.Ovoviviparous and apparently a uterine cannibal (oophagous), number of young 2 to 4 and rarely 6 in a litter off California(usually 2 to 4, commonly 4), and 3 to 7 in the eastern Atlantic. This species apparently uses inshore nursery areas intemperate waters (east coast of the United States, California, South Africa, the northeastern Atlantic and westernMediterranean, and probably elsewhere), with young sharks occurring in shallow bays (California, South Africa). In theeastern North Pacific (California) the species mates in summer, has a gestation period of nine months and gives birth duringthe spring. This shark matures between 3 and 8 years old, with a maximum age estimated at 45 to 50 years (Cailliet et al.,1983).Feeds mostly on small schooling fishes but also bottom fishes, including herring, sardines, shad, pilchards and menhaden(Clupeidae), anchovies (Engraulidae), lanternfishes (Myctophidae), lancetfishes (Alepisauridae), needlefishes (Belonidae),scad (Trachurus, Carangidae), mackerels (Scombridae), bluefishes (Pomatomidae), plaice and flounder (Pleuronectidae)and sole (Soleidae); also squids, octopi and pelagic crustaceans, and rarely seabirds. Herds and stuns its prey with its long,whip-like caudal fin, and is often caught on longlines by being tailhooked. It swims in narrowing circles around schools ofsmall fishes, splashing water with its caudal fin and compressing the school, then strikes and injures fishes with the caudal.Two threshers may cooperate on bunching and killing small fish.Size: The largest thresher. Maximum total length at least 573 cm and possibly to over 610 cm, with an estimated maximumat 651 cm from growth curves and older unconfirmed records up to 760 cm. Size at birth 114 to 160 cm, with term foetuses upto 139 to 156 cm and small freeliving specimens down to 117 to 145 cm. Immature males up to at least 252 cm, while anadolescent male examined was 288 cm and adult males are 314 to at least 420 cm. Females maturing at about 315 to400 cm, with immature or adolescent females up to 395 cm and adult females 376 to at least 549 cm. A length-weightequation is given by Kohler, Casey and Turner (1995) for fork length:W(kg) = 1.8821 x 10 -4 FL(cm) 2.5188(n = 88; both sexes)where: FL(cm) = 0.5474 x TL(cm) + 7.0262 (n = 13)

88 FAO Species Catalogue for Fishery Purposes No. 1Interest to Fisheries and Human Impact: Widely caught or formerly caught in offshore longline and pelagic gill netfisheries including those of the former USSR, Japan, Taiwan (Province of China), Spain, the USA, Brazil, Uruguay, Mexico,and other countries. Especially important areas for these fisheries are the northwestern Indian Ocean, the western, central,and eastern Pacific, and the North Atlantic. Also fished with anchored bottom and surface gill nets, and accidentally caughtin other gear including bottom trawls and fish traps. The species became the object of an important targeted pelagic gill netfishery off the west coast of the United States (particularly California but also Washington and Oregon) in the late 1970s,peaking at about 1 000 t in 1982 and declining due to overfishing to less than 300 t by the late 1980s. The targeted fisherywas ended by 1990 but the species was still caught as bycatch of the swordfish gill net fishery and may be sold for higherprices in the market than swordfish. The meat is highly prized fresh for human consumption (cooked) but is also eatensmoked and dried-salted; the fins are valuable for shark-fin soup; the hide is usable for leather and the liver oil can beprocessed for vitamins.Sports anglers seek these sharks in the USA, South Africa, and elsewhere with rod and reel. These sharks fight stronglywhen hooked and may jump out of the water. This and other threshers are listed as record fishes by the International GameFish Association.Apparently harmless to people, though the size of adults of this species should invite respect. There is an unconfirmedanecdotal account of a fisherman on the western North Atlantic coast of the USA that was decapitated by a tailstroke from abig adult thresher (Mundus and Wisner, 1971). A few assaults on boats are doubtfully attributed to this species. Smallspecimens have been seen underwater by divers, at the surface or close to the bottom, and have circled them at the limit ofvisibility without acting aggressively. Michael (1993) notes that this species is shy and difficult to approach underwater, butmentions an incident where a thresher of this species was aggressive toward a spearfishing diver off New Zealand. To thewriter’s knowledge, this species has seldom if ever been kept in captivity and is not currently the subject of ecotouristicshark-diving.The conservation status of this shark is little known but is of some concern despite its midrange intrinsic rebound potential (ameasure of the capacity to recover from fishing pressure; Smith, Au and Show, 1998) because of the history of the easternPacific thresher fishery (which declined quickly despite a relatively small and localized catch), and its exposure tohigh-intensity offshore fisheries virtually wherever it occurs.Local Names: Common thresher, Fox shark, Thrasher shark, Long-tailed shark, Sea fox, Sea ape, Slasher (England);Whiptail shark, Thintail thresher, Fynstert-sambokhaai (South Africa); Lluynog mor (Wales); Renard, Singe de mer, La faux,Poisson épée, Péi aspasu ratou, Touille à l’épée (France); Rabosa, Peix espasa, Zorra de mar, Pez zorro, Zorro blanco(Spain); Arequim, Peixe alecrim, Peixe raposo, Peixe zorra (Portugal); Peixe rato, Peixe cavallo (Madeira); Tubarâo raposo,Romano, Româo, Thresher shark (Azores); Pesce volpe, Pesce sorcio or Pavone, Pesce bandiera, Pesce pavone, Pescebannera, Volpe di mare, Pescio ratto, Allopia coda lunga, Allopia volpe marina, Pavone di mare, Pisce surci, Pisci cudalonga, Pisci cuduto, Pisci sciabula turca (Italy); Pas sabljas (Adriatic); Raefhajen (Sweden); Onagazame, Nadebuka,Nezumezame, Ma-onaga (Japan); Zorro (Cuba); Lisitska morskayia (Russia); Zorro cauda longa (Mozambique).Remarks: This account follows Bass, D’Aubrey and Kistnasamy (1975a) and Compagno (1984) in combining severalregional species of threshers from Chile, New Zealand and Australia into one wide-ranging species, A. vulpinus. Threshersexamined by the writer from the west coast of the USA and South Africa agree closely in morphology and meristics.Literature: Garman (1913); Bigelow and Schroeder (1948); Gubanov (1972, 1978); Bass, D’Aubrey and Kistnasamy(1975a); Hixon (1979); Gruber and Compagno (1981); Cailliet et al. (1983); Cailliet and Bedford (1983); Compagno (1984,1990b); Cailliet, Radtke and Welden (1986); Pascoe (1986); Bedford (1987); Cailliet and Radtke (1987); Moreno, Parajuaand Morón (1989); Cailliet, Holts and Bedford (1993); Hanan, Holts and Coan (1993); Michael (1993); Last and Stevens(1994); Kohler, Casey and Turner (1995); Eitner (1995); Smith, Au and Show (1998).2.2.6 Family CETORHINIDAEFamily: Subfamily Cetorhininae Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 397-398 (Family Lamnoidae).Emended to Family Cetorhinidae Gill, 1862, by Gill (1872, Smiths. Misc. Coll., [247]: 24).Type Genus: Subgenus Cetorhinus Blainville, 1816 (Genus Squalus Linnaeus, 1758).Number of Recognized Genera: 1.Synonyms: Group Selachina Günther, 1870 (Family Lamnidae): 389, 394. Emended to Family Selachidae Günther, 1870,by Poey, 1875: 85. Also Subfamily Selache (Family Lamna) Hasse, 1879: tab. 2. Type genus: Subgenus Selache Cuvier,1816 (Genus Squalus Linnaeus, 1758). Family Halsydridae Whitley, 1934: 196. Type genus: Halsydrus Neill, 1809.FAO Names: En - Basking sharks; Fr - Requins pélerin; Sp - Peregrinos.

Sharks of the World, Vol. 2 89Diagnostic Features: Head moderately long but much shorter than trunk. Snout moderately long, pointed and conical, notdepressed, flattened or blade-like. Eyes small with length 0.8 to 1.3% of precaudal length. Gill openings extremely large,width of first 17.7 to 29.2% of precaudal length, extending onto dorsal and ventral surfaces of head; all gill openings anteriorto pectoral-fin bases; gill rakers present on internal gill slits, in the form of hair-like modified dermal denticles with extremelyelongated crowns. Mouth large and arcuate, ventral on head; jaws hardly protrusable but greatly distensible laterally. Teethvery small, hook-like, not blade-shaped, and in 203 to 255/225 to 230 (432 to 480 total) rows. Several rows of small anteriorteeth in upper jaw, separated from the similar-sized laterals by a broad gap but without intermediate teeth; no symphysialteeth. Trunk fusiform and moderately stout, firm and not flabby. Caudal peduncle depressed and with strong lateral keelsand upper and lower crescentic precaudal pits. Dermal denticles large and rough, with erect hooked crowns, strong cuspsand ridges, with cusps of lateral denticles pointing in several directions. Pectoral fins long and moderately broad, muchshorter than head in adults; pectoral skeleton plesodic with radials extending far into fin webs. Pelvic fins smaller than firstdorsal fin but larger than second; fin skeleton aplesodic, not extending into fin web. First dorsal fin large, high, erect andangular; fin skeleton semiplesodic, extending partways into fin web. Second dorsal and anal fins moderately large but lessthan half size of first dorsal, with broad, non-pivotable bases. Caudal fin lunate, dorsal lobe moderately long but less thanone-third length of rest of shark, ventral lobe nearly as long as dorsal lobe. Neurocranium with a high cranial roof butotherwise low, not compressed, with long rostrum, depressed internasal septum and widespread nasal capsules, smallorbits with strong supraorbital crests, small stapedial fenestrae, and with hyomandibular facets not extended outward.Vertebral centra strongly calcified, with well-developed double cones and radii and prominent annuli. Total vertebral count109 to 116, precaudal count 50 to 54, diplospondylous caudal count 55 to 62. Intestinal valve of ring type with 47 to 51 turns.Size gigantic with adults 5 to about 10 m and possibly to 12 to 15 m long.Remarks: This family includes the single living genus Cetorhinus Blainville, 1816, and possibly a single living species, thebasking shark, C. maximus (Gunnerus, 1765) as well as several fossil species (Woodward, 1889; Cappetta, 1987).Publication of Cetorhinus Blainville and its synonym Selache Cuvier in the same year caused confusion, with severalprominent nineteenth century authors using Selache or the emended Selachus instead of Cetorhinus (Bonaparte, 1838;Müller and Henle, 1839; Swainson, 1838; Bleeker, 1859; Dumeril, 1865; Günther, 1870). Cetorhinus was used by severalnineteenth century authors (Gray, 1851; Gill, 1862b, 1872, 1893; Jordan and Gilbert, 1883; Jordan and Evermann, 1896)and was in general use by most twentieth century authors. Whitley (1934), followed by Fowler (1941) and Smith (1949),proposed that the genus Cetorhinus should be replaced by Halsydrus, which was based on the carcass of a ‘sea monster’washed ashore in the Orkney Islands and eventually identified as a basking shark. According to Bland and Swinney (1978),Halsydrus as originally proposed by Neill, 1809, was apparently a nomen nudum, separately proposed from descriptions ofthe Orkney ‘monster’, and does not comprise a senior synonym of Cetorhinus. Rafinesque’s (1810) genus Tetroras (and itsvariant Tetnoras) is hard to identify from its generic description, with Rafinesque’s claim that Tetroras has four gill openingsbeing probably erroneous (unless the specimen examined was abnormal). Rafinesque’s description of Tetroras angiova,the only species in the genus, does indeed suggest a basking shark in certain details (“denti in forma di raspa... ha gli occhipiccolissimi, e le aperture delle branchie bastantemente larghe.”). However, even if more evidence was available to provethat T. angiova actually was a basking shark, the substitution of Tetroras for Cetorhinus would not serve the stability ofzoological nomenclature due to universal usage of Cetorhinus for the basking shark at present.Couch (1862) proposed the genus Polyprosopus as separate from Cetorhinus and including two species apparently basedon aberrant basking sharks. Polyprosopus was recognized by Gill (1862b) but was synonymized with Cetorhinus orSelache by Dumeril (1865), Günther (1870) and subsequent authors.Following Müller and Henle (1839) and Bonaparte (1838, 1839), Cetorhinus or Selache were often placed in the family Lamnidae orIsuridae (Gray, 1851; Bleeker, 1859; Dumeril, 1865; Woodward, 1889; Regan, 1906a; Goodrich, 1909; Garman, 1913; Engelhardt,1913; Berg, 1940; Norman, 1966; Bailey et al., 1970; Nelson, 1976, 1984). Gill (1862b) proposed a subfamily Cetorhininae withinthe Lamnidae for Cetorhinus; the Cetorhininae as a subfamily of Lamnidae was recognized by Berg and Svidovidov (1955).Günther (1870) proposed a group Selachina within Lamnidae for Selache, which was followed by Hasse (1879). Gill (1872) elevatedCetorhininae to the family Cetorhinidae, which has been recognized by most modern authors (Jordan and Gilbert, 1883; Gill, 1893;Jordan and Evermann, 1896; Bridge, 1910; Jordan, 1923; Lozano y Rey, 1928; White, 1936, 1937; Bertin, 1939a; Romer, 1945,1966; Bigelow and Schroeder, 1948; Matsubara, 1955; Arambourg and Bertin, 1958; Glikman, 1964, 1967; Fowler, 1967a;Patterson, 1967; Blot, 1969; Budker and Whitehead, 1971; Lindberg, 1971; Rass and Lindberg, 1971; Pinchuk, 1972; Compagno,1973, 1981b, 1982, 1984, 1990b, 1999; Applegate et al., 1979; Chu and Meng, 1979; Gubanov, Kondyurin and Myagkov, 1986;Cappetta, 1987; Carroll, 1988; Eschmeyer, 1990, 1998; Robins et al., 1991a; Nelson, 1994; Helfman, Collette and Facey, 1997).Shirai (1996) in contrast placed Cetorhinus in a subfamily Cetorhininae of the family Alopiidae. Whitley (1940), Fowler (1941), andSmith (1949) substituted the family Halsydridae for Cetorhinidae after synonymizing Cetorhinus with Halsydrus (see above).Maisey (1985) synonymized the megamouth family Megachasmidae with Cetorhinidae, which was followed by Robins et al.(1991a). As noted by Springer and Gilbert (1976) and Compagno (1990b), Cetorhinus is very distinct. It may be the sister group ofthe family Lamnidae (Compagno, 1990b) but is apparently not closely related to Megachasmidae on its anatomy. Martin and Naylor(1997) suggested that Lamnidae and Cetorhinus are sister groups on gene similarities from cytochrome b DNA, while Long andWaggoner (1996) suggested that Megachasma and Cetorhinus are sister groups on dental morphology.Most authors recognize only one species of living basking shark (summarized in Garman, 1913; Bigelow and Schroeder, 1948;Springer and Gilbert, 1976; and Compagno, 1984), but Siccardi (1960, 1961) suggested that there are four species (or subspecies)of Cetorhinus, two from the North Atlantic and Mediterranean (C. maximus and C. rostratus), one from southern Australia (C.maccoyi) and one from the western South Atlantic (C. normani), based on morphometric, morphological and meristic differences.Characters included differences in head length, numbers of functional-tooth rows, gill opening width, body height, prepectoral

90 FAO Species Catalogue for Fishery Purposes No. 1length, trunk shape, interdorsal space, position of the dorsal and anal fins, vertebral numbers, and size. Apart fromArgentinean material of C. normani which she examined, Siccardi’s morphometrics were based on available literatureaccounts of basking sharks from other areas. More recently Tomés and Gomes (1989) suggested that two specimens ofBrazilian basking sharks they had examined differed morphologically from those from the North Atlantic.Springer and Gilbert (1976) suggested that there was insufficient evidence at present to separate the four Cetorhinusspecies on the basis suggested by Siccardi, with which Compagno (1984) concurred. The ‘small’ species C. rostratusseems to represent juveniles of C. maximus, and C. maximus, C. maccoyi, and C. normani as defined by Siccardi differedin characters that need more detailed analysis and consistent collection of data from basking sharks from various parts ofthe world to factor out individual variation, allometry, sexual dimorphism, and data idiosyncrasies. The question of allopatricspecies, subspecies, or populations within Cetorhinus needs further study. A problem with conventional morphologicalstudies is the lack of sufficient data for adequate comparisons, as well as the existence of data sets (includingmorphometrics and illustrations) that are difficult to compare. This stems in part from the great size of the basking shark,which makes data-collecting (particularly of morphometrics, as the writer can attest) and illustration difficult and prone toerror, as well as low and sporadic interest in basking shark variation, limited opportunities to collect data, and lack ofagreement on data protocols for morphometrics, meristics and descriptive morphology.Even if the basking shark comprises a single species, its known distribution suggests that there may be isolated populationsin temperate and boreal waters with potentially limited genetic interchange. Records of occasional basking sharks fromwarm-temperate to tropical inshore waters suggest, however, that interchange by crossing the tropics in deep water ispossible if infrequent. Populations or subpopulations of basking sharks could include a North Pacific population (withpossible eastern and western North Pacific subpopulations?), a North Atlantic population (with possible western NorthAtlantic, eastern North Atlantic and Mediterranean subpopulations?), a temperate South American population (with possibleeastern South Pacific and western South Atlantic subpopulations?), a southern African (Namibia and South Africa)population (‘Cape basking shark’), and an Australian-New Zealand population (if not forming separate New Zealand andAustralian subpopulations?). Some efforts are being made to collect tissue samples from Northern Hemisphere baskingsharks for DNA comparisons (UK CITES proposal, 1999), which should be extended to Southern Hemisphere sharks. Lackof recovery of basking shark numbers following intensive localized fisheries suggests isolated, geographically limitedpopulations or subpopulations that are not readily replaced through immigration. Some of these populations may be verysmall; basking sharks occurring off southern Africa and possibly Brazil and Argentina are apparently present in very smallnumbers compared to northern hemisphere localities.Cetorhinus Blainville, 1816Genus: Subgenus Cetorhinus Blainville, 1816 (Genus Squalus Linnaeus, 1758), Bull. Sci. Soc. Philomat. Paris, (8): 121.Type Species: not designated; Blainville included the species “Gunneri; Peregrinus; Shavianus; Homianus?” inCetorhinus without further comment. Gill (1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 398), designated Squalusmaximus “Linnaeus” (= Gmelin, 1788) as type of Cetorhinus, but this was not an included species. Jordan and Gilbert(1883, Bull. U. S. Nat. Mus., 16: 31) designated “Cetorhinus gunneri Blainv. = S. maximus L.” (a junior synonym of Squalusmaximus Gunnerus, 1765) as type of Cetorhinus, which may be the earliest valid type designation. Eschmeyer (1998,Cat.Fish.: CD-ROM) cited Jordan and Evermann (1896, Bull. U.S. Natn. Mus., (47), pt. 1: 51) as a later, similar type designation.Number of Recognized Species: 1.Synonyms: Genus Halsydrus Neill, 1809a: 5-6; also Neill, 1809b: 90-91 (nomina nuda, not seen; Bland and Swinney,1978: 134; Eschmeyer, 1998: CD-ROM). ?Genus Tetroras Rafinesque, 1810a: 11. Type species, Tetroras angiovaRafinesque, 1810, by monotypy. Also Rafinesque, 1810b: 46; Rafinesque, 1815: 93. Subgenus Selache Cuvier, 1816(Genus Squalus Linnaeus, 1758): 129. Type species, “Sq. maximus L.” by monotypy (Squalus maximus Gmelin, 1788, ajunior synonym of Squalus maximus Gunnerus, 1765). Genus Halsydrus Fleming, 1817: 713; Fleming, 1822: 380. Typespecies: Halsydrus pontoppidani Fleming, 1817, by monotypy? (Bland and Swinney, 1978: 134). ?Genus ScoliophisAnonymous, 1817: 44, cited by Sherborn (1930: 5809) and Neave (1940: 150). Listed in synonymy of Halsydrus maximusby Whitley (1939: 241) without explanation; possibly a ‘sea serpent’ based on a basking shark carcass? Genus SelancheJarocki, 1822: 452 (error for Selache Cuvier, 1816?). Genus Selachus Minding, 1832: 52 (unjustified emendation of SelacheCuvier, 1816?). Genus Ceteorhinus Agassiz, 1846: 75 (error or emendation for Cetorhinus Blainville, 1816?). GenusPolyprosopus Couch, 1862: 67. Two species, P. rashleighanus (Couch, 1832) and P. macer Couch, 1862 included in thegenus without a type designation. Type species, “Polyprosopus Rashleighanus Couch”, by subsequent designation of Gill(1862b: 398) and according to Eschmeyer (1998: CD-ROM). Genus Hannovera van Beneden, 1871: 504. Type species:Hannovera aurata van Beneden, 1871, possibly by original designation. Based on fossil basking shark gill rakers from thePliocene of Belgium (Woodward, 1889), possibly the same species as the living basking shark (Cappetta, 1987). GenusCethorhinus Escribano, 1909: 340 (error or emendation for Cetorhinus Blainville, 1816?). Genus Scapasaurus Marwick,1942: 1. Nomen nudum, no species name, apparently proposed for a ‘sea monster’, based on the carcass of a baskingshark washed up in Scapa Flow. The vertebrae of this shark are in the British Museum (Natural History), BMNH 1946.8.3.1(citation from Bland and Swinney, 1978: 134). Genus Tetnoras Fowler, 1941: 112. Based on Tetroras Rafinesque, 1815: 93,the printing of which resembles Tetnoras in the original but seems to be a poorly printed “r” rather than an “n”. ?GenusTetraoras Bigelow and Schroeder, 1948: 146. Variant spelling on Tetroras Rafinesque, 1810? Genus Hannoveria Capetta,1987: 107. Variant spelling on Hannovera van Beneden, 1871.click for next page

click for previous pageSharks of the World, Vol. 2 91Cetorhinus maximus (Gunnerus, 1765) Fig. 67Squalus maximus Gunnerus, 1765, K. Norske Vidensk-selsk.Scr. Trondh.: 33, pl. 2. Holotype, apparently none. TypeLocality: Trondhjem, Norway.Synonyms: Halsydrus pontoppidani Neill, 1809: 5-6; also Neill, 1809: 90-91 (nomina nuda; Bland and Swinney, 1978:134). ?Tetroras angiova Rafinesque, 1810a: 11. No types?, species dubium. Also Rafinesque, 1810b: 46; Rafinesque,1815: 93. Squalus gunnerianus Blainville, 1810: 256, pl. 2, fig. 3; also Blainville, 1810b: 170. Types? (Eschmeyer, 1998:CD-ROM). Squalus homianus Blainville, 1810: 257, pl. 2, fig. 1. Also Blainville, 1816: 121. Types? (Eschmeyer, 1998:CD-ROM). Squalus pelegrinus Blainville, 1810a: 256, pl. 2, fig. 2. Holotype: Museum National d’Histoire Naturelle, Paris,MNHN 9853 (Eschmeyer, 1998: CD-ROM). Squalus peregrinus Blainville, 1811: 365. Also Blainville, 1816: 121. Variant andpossibly emended spelling of Squalus pelegrinus Blainville, 1810. Squalus (Cetorhinus) Gunneri Blainville, 1816: 121(nomen nudum). Squalus (Cetorhinus) Shavianus Blainville, 1816: 121 (nomen nudum). ?Scoliophis atlanticus Anon.,1817: 44, cited by Sherborn (1923: 535). Listed in synonymy of Halsydrus maximus by Whitley (1939: 241) withoutexplanation, possibly a ‘sea serpent’ based on a basking shark carcass? Halsydrus pontoppidani Fleming, 1817: 713.Stronsa, Orkney Islands. Holotype: National Museum of Scotland, NMSZ-1979.012, three vertebrae in alcohol and driedceratotrichia, fide Herman, McGowan and Swinney (1990: 1). Squalus isodus Macri, 1819: 76, pl. 1, fig. 2. Mediterranean.No types (Eschmeyer, 1998: CD-ROM). Squalus rostratus Macri, 1819: 76, pl. 1, fig. 1, pl. 2. Mediterranean. No typesknown (Eschmeyer, 1998: CD-ROM). Squalus elephas Le Sueur, 1822: 350, pl. Types: Based on a large adult malespecimen, about 10 m (32 ft 10 in) TL from the New Jersey coast, possibly not saved. Types? (Eschmeyer, 1998: CD-ROM).Squalus rashleighanus Couch, 1838: 51. Type locality, Cornwall. Types? (Eschmeyer, 1998: CD-ROM). Squalusrhinoceros Mitchell, in DeKay, 1842: 358 (reference to name in newspaper article by Mitchell, 1828 citation unknown).Name only, not available, Maine Coast (Eschmeyer, 1998: CD-ROM). Squalus cetaceus Gronow, 1854: 6. Type locality,Norway. No types known (Eschmeyer, 1998: CD-ROM). Polyprosopus macer Couch, 1862: 68, pl. 15, fig. 2. Type locality,Startpoint, Cornwall, England. No types known (Eschmeyer, 1998: CD-ROM). Cetorhinus blainvillei Capello, 1870: 233, 1pl. Type locality, Portugal. Types? (Eschmeyer, 1998: CD-ROM). Selachus pennantii Cornish, 1885: 351. Type locality:Cornwall. No types known (Eschmeyer, 1998: CD-ROM). Tetroras maccoyi Barrett, 1933: 13. Types? Australia. Referencefrom Whitley (1934: 197) and Eschmeyer (1998: CD-ROM). ?Tetraoras angiona Bigelow and Schroeder, 1948: 146. Errorfor Tetroras angiova Rafinesque, 1810. Cetorhinus maximus forma infanuncula van Deinse and Adriani, 1953: 309.Holland, for specimens without gill rakers. Types? Not available, according to Eschmeyer (1998: CD-ROM). Cetorhinusmaximus normani Siccardi, 1960: 255, pl. 1. Syntypes (4): Museo Argentino de Ciencias Naturales, Bernardino Rivadavia,Capital Federal, Argentina, MACN (Eschmeyer, 1998: CD-ROM). Also Siccardi, 1961: 96-97 (as species, C. normani).Other Combinations: Halsydrus maximus (Gunnerus, 1765), Selache maxima, Selache maximus or Selache maximum(Gunnerus, 1765), Selache elephas (Le Sueur, 1822).FAO Names: En - Basking shark; Fr - Pélerin; Sp - Peregrino.Fig. 67 Cetorhinus maximusField Marks: The great size, enormous gill slits that virtually encircle the head, dermal denticle gillrakers, pointed snout, huge, subterminal mouth with minute hooked teeth, caudal peduncle withstrong lateral keels, and lunate caudal fin distinguish this shark from all others. Colour: blackish togrey-brown, grey or blue-grey above and below on body and fins, undersurface sometimes lighter,often with irregular white blotches on the underside of the head and abdomen; flanks sometimes withlighter linear striping and spots.Diagnostic Features: See family Cetorhinidae above.UNDERSIDE OF HEAD

92 FAO Species Catalogue for Fishery Purposes No. 1Distribution: Circumglobal with a wide but possibly disjunct distribution: Western North Atlantic: Canada (Newfoundland,Nova Scotia, New Brunswick) and the United States (Maine, New Hampshire, Massachusetts, Connecticut, Rhode Island,New York, New Jersey, Delaware, Maryland, Virginia, North Carolina, Georgia and Florida, also northern Gulf of Mexico),with most records in the Gulf of Maine and in the Mid-Atlantic Bight. Western South Atlantic: Southern Brazil, Uruguay andnorthern Argentina, Falkland (Malvinas) Islands. Eastern North Atlantic, Arctic Ocean, and Mediterranean Sea: Iceland(south and west coast), Faeroe Islands, Norway and Russia (western Barents Sea and White Sea), south to Sweden,Scotland, Ireland, England, France, Spain, and Portugal, the entire Mediterranean Sea (including Italy, Greece and Turkey),Morocco, and Senegal. Eastern South Atlantic and southwestern Indian Ocean: Namibia and South Africa (Northern andWestern Cape Provinces). Western North Pacific: Japan, Korean Peninsula, Taiwan (Province of China), China. WesternSouth Pacific: Australia (New South Wales, Victoria, Tasmania, South Australia and Western Australia), New Zealand.North-central Pacific: Hawaiian Islands (USA). Eastern North Pacific: Gulf of Alaska and Aleutian Islands to Gulf ofCalifornia, including Canada (British Columbia), the United States (Alaska, Washington, Oregon, and California), andMexico (Baja California and northern Gulf of California). Eastern South Pacific: Ecuador, Peru and Chile, GalapagosIslands.Habitat: A coastal-pelagic and semioceanic or oceanic shark found in boreal to warm-temperate waters of the continentaland insular shelves, occurring well offshore and often very close to land, just off the surf zone, and entering enclosed bays.Most individuals are observed in shallow coastal waters, but records from aerial cetacean and sea turtle surveys in thewestern North Atlantic, a stranding at an oceanic island chain (Hawaiian Islands), and pelagic driftnet records in thetemperate North Pacific suggests that it readily ventures into the epipelagic zone. It has been sighted at the surface over theslopes from 200 to 2 000 m, and with a few sighted in the oceanic basins at 2 000 to 4 000 m. The basking shark is thought tooverwinter in deep offshore waters, possibly near the bottom, and has occasionally been caught in deep bottom trawls in theGulf of St. Lawrence and off Scotland. Off the United Kingdom, Japan and Newfoundland most records are in waters of 8 to14°C, though off the Atlantic coast of New England sightings were in waters of 11 to 24°C with most at 16 to 24°C. Baskingsharks seem to prefer ocean fronts, where differing water masses meet, and close inshore off headlands, around islandsand in bays with strong fluctuation of water masses from tidal flow and where aggregations of zooplankton occur.Biology: The sociobiology, behaviour and population biology of the basking shark are sketchily known and need furtherinvestigation. Although basking sharks are huge impressive animals that are readily accessible to underwater observationand are conspicuous to surface observers, a limited amount is known of displays, courtship, and social structure, though thisis changing with intensive behavioural observation of basking sharks off the UK (Earll et al., 1992). A problem is that thebasking shark is traditionally studied and viewed as a fisheries object, so that many details of its behaviour and generalbiology remain to be elucidated.Basking sharks are apparently social animals as in some other lamnoids, and show some behaviours that may beintraspecific displays. This shark is often seen at or near the surface, singly, in pairs or triads or in schools up to a hundred ormore individuals, basking with dorsal fins out of the water or even with bellies upward. Adults have been seen together, ashave pairs with young individuals and adult females (the former assumed to be offspring). These sharks are often seenmoving slowly forward or in short arcs with their mouths distended like hoops and pharynx expanded while feeding,sometimes with snouts out of the water. They also cruise when not feeding, with mouth partly closed and pharynx notexpanded. The open-mouthed feeding posture of basking sharks is analogous to gaping in certain other lamnoids, although

Sharks of the World, Vol. 2 93it is not established if the basking shark uses gaping in a social context apart from feeding. Surface-basking in this shark isthought to be correlated with surface concentrations of zooplankton and also with courtship and mating. Two, three or moreindividuals may engage in tandem-swimming, in a straight line or in circles, which suggests to some writers that a row ofthese sharks swimming together may have been mistaken for a single huge ‘sea serpent’ in the past. They also showparallel-swimming or stalling (stopping, sometimes for protracted periods), with sharks alongside one another, andsometimes make a wedge-formation of three individuals while feeding. Nose to tail circling has been described with a maleand female and even a pair of males. The sharks direct their feeding along tidal lines, may circle zooplankton patches, andmay swim in one direction, do a U-turn, then parallel their previous track. They can also stop and gulp down planktonconcentrations, may stall and keep in one position, may swim very slowly, and show relatively fast dashes at the surface.They may dive suddenly when disturbed, and may flick their tails sharply before doing so. Individuals may droop their dorsalfins nearly 45° to one side; they also may lift their tails out of the water and do a tail-sweep, wagging the tail from side to sideat a 45° angle in an exaggerated movement. Under provocation (particularly when harpooned) this species may defenditself by tail-swatting. Basking sharks may approach divers and boats quite closely, possibly out of curiosity, and swimaround them or follow them.Recent observations and photographs confirm that basking sharks breach or jump partway or entirely out of the water, and ithas been suggested that they do so to dislodge parasites or commensals such as lampreys or remoras. In addition to theectoparasitic copepods found on other sharks, basking sharks often have sea lampreys (Petromyzon marinus) attached totheir skin in the North Atlantic, and although lampreys apparently are unable to cut through the formidable denticle-armouredskin of the shark, they may be enough of an irritant to evoke a reaction like breaching or rubbing on objects or the bottom(chafing) to dislodge them. Basking sharks may breach alone or while in groups, and breaching may also have a socialsignificance in intraspecific communication.Dead basking sharks are often stranded on the beach, in a state of advanced decay and damaged and partly dismemberedfrom rolling in the surf, and they have been also occasionally lifted from the bottom by trawls on fishing boats. Several timessuch carcasses have been misidentified as ‘sea serpents’, plesiosaurs or other fabulous monsters, some of which havebeen given specific and even generic names.Numbers of basking sharks sighted may fluctuate greatly in given areas each year, with irregular increases (‘invasions’) anddecreases that are of uncertain cause. Estimates of world population numbers are unavailable, as with most other sharks,but aerial surveys suggest that numbers of basking sharks in localized areas (e.g., New England coast and Monterey Bay)may not exceed 2 000 to 6 000, nor have catches in the eastern North Atlantic much exceeded 3 000 per year when catcheswere highest.The massive liver of the basking shark, contained in a long body cavity, apparently serves as a ‘hepatic float’ to adjust it toapproximately neutral buoyancy.Basking sharks are highly migratory, and noteworthy for their seasonal appearance in numbers in given localities andsubsequent disappearance. Off the Atlantic seaboard of North America they appear in the southern part of their range inspring (North Carolina to New York), apparently shift northward in summer (New England and Canada), and disappear inautumn and winter. Off the eastern North Pacific basking sharks occur in greatest numbers during autumn and winter in thesouthernmost part of their range there (California), but shift at least in part to more northern latitudes in spring and summer(up to Washington and British Columbia). Off the United Kingdom the bulk of the population there appears in the springtimeand disappears by autumn, but individuals may be present at all seasons of the year. Aerial surveys off New Englandsuggests that basking sharks move into shallow coastal waters from deeper water in springtime as zooplankton bloomsdevelop; sightings there indicate that basking sharks occur at the surface in the epipelagic zone in deep water, above theslopes and even in the ocean basins. Research is currently in progress in the eastern North Atlantic using satellite radio tagsto resolve some of the problems of migration in this species. Priede (1984) details an early successful short-term satellitetracking effort on a radio-tagged basking shark, and plans for satellite tagging using more modern equipment are underwayin the United Kingdom (Dunbar, Fowler and Denham, 1995). This space-age methodology may unravel some of the secretsof the basking shark’s seasonal migrations and movements. Adult individuals found off the United Kingdom in summertimeare apparently engaged in courtship activity and copulation, as indicated by behavioural observations and courtship andmating scars found on captured individuals.Pronounced spatial and seasonal populational segregation may be a characteristic of this species, as suggested byfisheries catches off the United Kingdom. Most individuals caught there in the summer were subadult or nonpregnant adultfemales, outnumbering the males by 40:1, but in the winter the few individuals caught were mostly males. Off Japan mostbasking sharks taken in fisheries are or were females.Pregnant females are almost entirely unknown for the species, suggesting that such females are spatially andbathymetrically separated from those members of the population that are regularly seen basking at the surface. It is alsopossible that the number of pregnant females is very low relative to the number of non-pregnant adult females, withcorrespondingly few being taken and with extremely low fecundity for the species. Juveniles below 3 m long have beenextremely rare in the literature, with a single record of a freeliving individual about 1.7 m long reported from the British Isles.More recently, field observers from the UK Marine Conservation Society have reported larger numbers of sightings ofindividuals between 2 and 4 m long (34%), 4 to 6 m (about 38%), and 6 to 8 m (about 18%), but with few young below 2 m(3%) or large adults above 8 m (8%) of over 3 300 individuals sized (Pollard, 1996). This suggests low local recruitment ofyoung, but large numbers of juveniles (2 to 6 m) and smaller numbers of adults (6 to 8+ m).

94 FAO Species Catalogue for Fishery Purposes No. 1A pregnant female has been reported with a litter of six young. Adult, nonpregnant female basking sharks have immensenumbers of small eggs in their ovaries, which suggests that this shark has uterine cannibalism as in many other lamnoids,with embryos feeding on the small eggs (oophagy). Feeding on smaller siblings (adelphophagy) has not beendemonstrated. Young below 2 m appear in waters of the United Kingdom in springtime, suggesting spring pupping.Age of this shark has been estimated by counting vertebral rings and attempting to correlate them with supposed changes insize of individuals within a population, but this has met with limited success. It has been suggested that birth occurs after a3.5 year gestation period, and that two calcified rings per year are laid down until maturity at between 6 and 8 years for males(Parker and Stott, 1965). The correlation of the rings with time is uncertain and controversial. Pauly (1978) initially suggesteda yearly rate of ring deposition, but Pauly (in press) recently suggested that vertebral rings were unreliable for estimates ofage in basking sharks due to variation in numbers of rings along the vertebral column and apparent lack of correlation ofvertebral ring numbers to yearly cycles. Pauly (in press) used size frequency data to develop a von Bertalanffy growth curvefor the basking shark, assuming a size at birth of 1.5 m (see below), a gestation period of about 2.6 years, and an age atgreatest length (10 m) of about 50 years. There may be a possible cycle of two to three years between litters, and a possibleage at maturity for males at 12 to 16 or more years and up to 20 years for females (UK CITES Authority, 1999).The basking shark is one of the four types of large, filter-feeding elasmobranchs, the others being the megamouth and whalesharks, and the manta rays (Mobulidae). The basking shark may be unique in relying entirely on the passive flow of waterthrough its pharynx generated by swimming for filtration; the other filter-feeders may assist the process of food ingestion byactively pumping or gulping water and food organisms into their pharynxes. The basking shark feeds on small planktonicorganisms trapped on its unique gill rakers, apparently with the help of mucus secreted in its pharynx. Food items includesmall copepods (including calanids), barnacle, decapod, stomatopod larvae and fish eggs. On the average a half tonne ofmaterial may be present in the stomach of these sharks. While feeding the basking shark usually cruises with mouth widelyopen and gills distended, occasionally closing its mouth to ingest its prey. An average adult has been estimated to becapable of filtering over 2 000 t of water per hour assuming a constant cruising speed of about 2 knots. They seem to feedalmost at random in low plankton densities, but are highly selective and actively search in restricted areas for high densitiesof zooplankton above a minimum concentration before feeding. Off southwestern England they actively seek planktonconcentrations at the surface along boundaries of tidally influenced water masses or tidal fronts over bottom about 20 to50 m deep.The facts that the basking shark periodically sheds its gill rakers and that plankton densities seasonally fall below levelsthought essential to maintain ordinary swimming and metabolic activity in this shark have spawned a controversy overwhether or not the species remains active when deprived of gill rakers and high plankton densities. It has been suggestedthat the basking shark may hibernate on the bottom, perhaps at the edges of continental shelves or on the slopes, until itsrakers are replaced and plankton blooms reoccur. Proof of hibernation has never been forthcoming, and an alternatehypothesis has been suggested that the basking shark may turn to benthic feeding when it loses its gill rakers. A possibleadditional factor is that the massive, oil-filled liver of this species may serve as a metabolic store to supply energy to supporta reduced rate of activity (slower swimming in colder, deep water) while gill rakers regenerate and plankton supplieseventually recuperate. Estimates have been proposed that, in north European waters, the basking shark drops its gill rakersin early winter and takes about 4 or 5 months to fully replace them. An anecdotal report suggests that basking sharks mayfeed on small schooling fishes such as herring as a possible alternative to plankton-sieving, but this remains to be confirmed.By far the most important enemy of this shark is humanity, primarily from fisheries but also from collisions with boats. A deadbasking shark with wounds was once seen near a pod of killer whales (Orcinus orca), with the implication that the whaleskilled the shark. White sharks (Carcharodon carcharias) will readily consume meat from dead basking sharks, but have notbeen recorded as attacking living ones. A large white shark or a group of white sharks are sufficiently formidable to kill abasking shark. Lampreys, copepods and cookiecutter sharks (Isistius) may produce external wounds, but it is not known ifthey contribute to mortality.Size: Basking sharks have been credited as reaching a maximum total length of 12.2 to 15.2 m, but even if this is correctmost specimens do not exceed approximately 9.8 m. Pauly (in press), analyzing the largest North Atlantic records of baskingsharks in an ‘extreme value theory plot’, suggested that individuals over 10 m long were unlikely to exist. This is supported byobservational data off the UK (Pollard, 1996), in which few basking sharks (8% of over 3 300) over 8 m long were seen. Sizeat birth is uncertain with few pregnant females examined. An unconfirmed record of a foetus about 1.7 m long, anotherfoetus 1.5 m long, and a 165 cm freeliving individual suggests that size at birth may be about 1.5 to 1.7 m, and hence isslightly greater than any other known ovoviviparous or viviparous shark except perhaps the common thresher and whiteshark. Males mature between 4 and 7 m and reach about 9 m; a late adolescent male 6.85 m long from Cape Town, SouthAfrica (dissected by the writer) had almost fully calcified claspers; six adult males examined by Matthews and Parker (1950),Springer and Gilbert (1976), and the writer were 7.01 to 8.46 m long (average 7.83 m). Females are immature at 3.19 and3.59 m (Springer and Gilbert, 1976, L.J.V. Compagno, unpub. data), females are mature at 8.0 to 9.8 m; six adult femalesexamined by Matthews and Parker (1950) and Springer and Gilbert (1976) were 8.03 to 8.48 m long (average 8.27 m),suggesting that females slightly exceed males in size as with many other sharks. Pauly (in press) suggested a maximumweight of 7.5 t at 10 m using an isometric weight curve, W(t) = 0.0075 * TL(m) 3 . The basking shark is apparently the secondlargest shark, neoselachian elasmobranch, and living fish-like vertebrate after the whale shark (Rhincodon typus), althoughit may be rivalled in size by the manta (Manta birostris).

Sharks of the World, Vol. 2 95Interest to Fisheries and Human Impact: The basking shark has been the object of small-scale targeted harpoon fisheriesfrom small boats and small ships (some built and equipped like whale-catchers) during the eighteenth, nineteenth andtwentieth centuries. In the twentieth century such fisheries operated primarily off the Norwegian coast, Ireland, Scotland,Iceland, Spain, United States (California), Peru, Ecuador, China and Japan. It has been sporadically if heavily fished due toperiodic depletion of basking shark stocks or intrinsic factors of the fisheries such as falling demand for this shark. During theeighteenth and nineteenth centuries basking sharks were also harpooned from large whaling vessels in the Gulf of Maine(United States and Canada) and probably elsewhere; basking sharks in the western North Atlantic were apparentlydecimated by such activities. The basking shark has also been taken as discarded and utilized bycatch in net gear, includingbottom anchored gill nets, floating gill nets, pelagic gill nets, bottom trawls, pelagic trawls, and even anchored fish weirs. It iscaught as a rare bycatch of gill netters targeting thresher sharks and swordfish off California and was taken by squid gillnetters as an uncommon bycatch in the epipelagic zone of the temperate North Pacific. It was formerly perceived as aproblem to salmon gill netters off British Columbia and elsewhere in the Pacific Northwest by fouling and damaging gill nets,and was formerly subjected to a localized eradication programme in British Columbia using a modified fisheries vessel with aknife-like ram on its bow to impale and kill sharks at the surface. The activities of this vessel in the 1950s apparentlydecimated the basking shark locally, which has not shown signs of recovery almost fifty years later.Meat from the basking shark is used fresh or dried-salted for human consumption. The fins of the basking shark are used forshark-fin soup and currently are extremely valuable, with the huge pectoral and dorsal fins recently (1999) sold forUS$10 000 to 20 000 each. The basking shark was traditionally targeted for its liver, which is rich in oil and very large. Theliver oil of the basking shark was formerly used for tanning leather and for lamp oil; it also yields vitamin A, and in moderntimes has been extracted for its high level of squalene which is used for medicinals and cosmetics. The hide of the baskingshark is processed for leather, the cartilage for medicinals, and the carcass is sometimes rendered into fishmeal. Baskingshark fisheries have not been monitored in detail except in the eastern North Atlantic and New Zealand, and catch statisticsfor basking sharks have been reported to FAO from New Zealand, Portugal, France and Norway. Norwegian catches werethe highest reported for any nation, with catches of 2 200 to 18 700 t reported to FAO in the 1960s and 1970s, but havedeclined to less than 500 t in 1997 and have been caught under a quota from 1978 onward. Slightly more than 12 000 sharkswere caught off the west coast of Ireland over a 29 year period, with peak catches in the 1950s and apparently no recovery inrecent years. New Zealand, Portugal and France report tiny catches of 1 to 14 t at present, with the New Zealand catchstrictly regulated as utilized bycatch only and with targeted fisheries not allowed.The basking shark is usually quite tolerant of boats approaching it, which makes it easy to hunt with harpoons from smallboats. Divers have been able to swim up to individuals and photograph them without invoking flight reaction. Basking sharksmay approach divers quite closely, possibly out of curiosity, and swim around them. This species is regarded as ordinarilyharmless and inoffensive when not provoked or molested. The immense size and power of the basking shark should inviterespect by ecotouristic divers who swim with these sharks. Divers should take care in contacting the skin of basking sharks,which has large dermal denticles with sharp, hooked crowns that point forward and sideways as well as backward. As thewriter can attest from dissecting large basking sharks, these denticles can inflict irritating lacerations on unprotected skin.There is considerable and growing concern over the conservation status of the basking shark because of declining numberscaught over the last five decades, with total catches dropping from 0.5 to 0.2 of their peaks during the 1970s and with somelocalized fisheries depressed to less than 0.1 of their peaks in 10 to 25 years (UK CITES , 1999). Localized basking sharkfisheries have a short-term boom and bust nature, and apparently are not sustainable at even moderate levels for very longdue to the biological limitations of the basking shark as a fisheries species. The basking shark has proved to be extremelyvulnerable to overfishing, perhaps more so than most sharks, and this can be ascribed to its slow growth rate, late maturity,long gestation period, probably low fecundity, long life, few predators when adult, probable small size of existing populations(belied by the immense size of individuals in their small schools), ready access to small fishing vessels inshore and offbuilt-up areas, and possibly little interchange between populations or stocks. A recent and major problem is theextraordinarily high value of basking shark fins, which promotes finning of sharks caught as bycatch of other fisheries ratherthan the release of captured individuals alive. The fin trade also supports small-scale targeted fisheries beyond thecommercial limits imposed by the lesser value of other basking shark products such as liver oil, and by the depleted stocks ofbasking sharks in most parts of the world. High fin prices can also promote illicit poaching and illegal trade despite local,national and international protective measures.The basking shark has been placed on the IUCN Red List as a vulnerable species, and has received protection fromexploitation in the territorial waters of the United Kingdom, Malta, New Zealand and the United States (East Coast, includingseparate protection by the State of Florida in its territorial waters). It was sponsored for CITES listing on Appendix II by theUnited Kingdom (for the 2000 CITES meeting), to promote regulation of basking shark catches and limits to internationaltrade in fins and other products. However the CITES proposal was very narrowly defeated and the UK is going on to proposethe basking shark for CITES Appendix III listing, effective from 2000 onward. The basking shark will be fully protected in theentire Mediterranean Sea once the Barcelona Convention for the Protection of the Mediterranean Sea is ratified by itsMember Nations. It is listed as a strictly protected species of the Bern Convention on Conservation of European Wildlife. It isunder consideration for total protection off South Africa (2000).The basking shark has become a ‘supershark’, with a high-profile and fortunately positive public image, and has a strongand highly motivated conservationist following in the United Kingdom. This is somewhat similar to, albeit better organizedthan conservationists promoting the whale shark in the western Atlantic and the Indo-Pacific, which ensures publicawareness of the conservation problems and interest in protecting both ‘gentle giants’. The basking shark is being monitoredby three different conservation groups in the United Kingdom, with sightings by volunteers reported on standard data-cards.

96 FAO Species Catalogue for Fishery Purposes No. 1Local Names: Basking shark, Giant basking shark, Elephant shark, Capidolo, Oilfish, Pelerin, Sun fish or Sunfish, Sail fishor Sailfish, Hoe mother or Homer, Bone shark, Gurry shark (English); Hoe mother, Homer (Orkneys); Heulgi (Wales);Cearban (Scotland); Sun fish, Liabhán mór, Liabhán chor gréine (Ireland); Haar moer (Scandinavia); Brugde, Brygde, Rybrigde (Norway); Brugden (Sweden); Rymer, Beinhaakal (Iceland); Brugde (Denmark); Mandelhai, Riesenhai (Germany);Reusenhaai (Belgium); Reuzenhaai (Holland); Le pélerin, Le très grand, Le squale a fanons, Poisson à volies, Elephant demer (France); Squalo massimo, Cagnea, Selachio gigante, Cagnia, Imbestinu, Caniscu, Squalo elefante (Italy); Pixi tunnu(Malta); Carago, Peixe carago, Peixe frade (Portugal); Peregrino (Spain); Peixe frade, Tubarâo frade, Basking shark(Azores); Psina golema (Adriatic); Ubazame or Old woman shark, Teguzame [Tenguzame?] or Long-nosed shark,Bakazame or Foolish shark, Zozame or Elephant shark (Japan); Elefante (Cuba); Peje-vaca or Cow-fish (Chile); Akulagigantakaia, Akuloobraznye, Bol’shezhabernye akuly, Gigantskie akuly, Akuly nastoiaschie, Lao sha k’o (Russia); Southernbasking shark (Australia); Cape basking shark, Koesterhaai (South Africa).Literature: Garman (1913); Barnard (1925, 1937, 1947); Lahille (1928); Fang and Wang (1932); Whitley (1934, 1939, 1940,1967); Norman (1937); Fowler (1941, 1967a); Hildebrand (1946); Bigelow and Schroeder (1948); Matthews (1950, 1956);Matthews and Parker (1950a, b); Van Deinse and Adriani (1953); Parker and Boeseman (1954); Lindberg and Legeza(1959); Siccardi (1960, 1961); Baldwin (1961); Larkins (1964); Parker and Stott (1965); Kato, Springer and Wagner (1967);Squire (1967, 1990); Lindberg (1971); Shiino (1972, 1976); Miller and Lea (1972); Hart (1973); Sadowsky (1973); Antezana(1977); Bass, D’Aubrey and Kistnasamy (1975a); Springer and Gilbert (1976); Pauly (1978, in press); Applegate et al.(1979); Konstantinov and Nizovtsev (1979); Davis (1983); Silva-Santos, Gomes and Ferreira (1983); Compagno (1984,1990a, b); Nakaya (1984); Quero (1984); Kenney, Owen and Winn (1985); Horsman (1987); Paulin et al. (1989); Tomas andGomes (1989); Herman, McGowan and Swinney (1990); Springer (1990); Earll et al. (1992); Hanan, Holts and Coan (1993);Izawa and Shibata (1993); Michael (1993); Bonfil (1994); Darling and Keogh (1994); Last and Stevens (1994); Dunbar,Fowler and Denham (1995); Taylor (1995); Uchida (1995); Pollard (1996); Fowler (1996); Santos, Porteiro and Barreiros(1997); Fairfax (1998); McEachran and Fechhelm (1998); Sims and Quayle (1998); Castro, Woodley and Brudek (1999); UKCITES Authority (1999); D. Ebert (pers. comm., on Namibian records); S. Fowler (pers. comm.); J. Naughton (pers. comm.,on Hawaiian record); L.J.V. Compagno (unpub. data from South Africa).2.2.7 Family LAMNIDAEFamily: Lamnoidea Müller and Henle, 1838a, Mag. Nat. Hist., new ser., 2: 36. Also Subfamily Lamnini Bonaparte, 1838,Nuov. Ann. Sci. Nat., Bologna, ser., 1, 2: 209 (Family Squalidae); Family Lamna Hasse, 1879, Nat. Syst. Elasmobr., (1): 52.Emended to Family Lamnidae Müller and Henle, 1838 by Richardson, 1846, Ichthyol. China Japan: 195.Type Genus: Lamna Cuvier, 1816.Number of Recognized Genera: 3.Synonyms: Tribe Isurina Gray, 1851 (Family Squalidae): 58. Emended and raised in rank to Family Isuridae Gray, 1851 byGill, 1893: 130. Type genus: Isurus Rafinesque, 1810. Subfamily Carcharodontinae Gill, 1893 (Family Isuridae): 130. Typegenus: Carcharodon Smith, 1838. Family Carcharodontidae Whitley, 1940: 68. Independently proposed from Gill, 1893.Type genus Carcharodon Smith, 1838. Family Lamiostomatidae Glikman, 1964: 11, 105. Type genus LamiostomaGlikman, 1964: 105.FAO Names: En - Mackerel sharks, Porbeagles, White sharks; Fr - Requins taupe; Sp - Jaquetones, Marrajos.Field Marks: Large sharks with pointed snouts and spindle-shaped bodies, long mouths with large blade-like teeth, long gillslits, long pectoral fins and high first dorsal fins, small pivoting second dorsal and anal fins, large lateral keels and prominentprecaudal pits on the caudal peduncle, and lunate caudal fins.Diagnostic Features: Head moderately long but shorter than trunk. Snout moderately long pointed and conical not greatlyelongated flattened or blade-like. Eyes small to moderately large, length 0.9 to 3.4% of precaudal length. Gill openings large,width of first 7.6 to 10.8% of precaudal length, extending onto dorsal surface of head; all gill openings anterior to pectoral finbases; no gill rakers on internal gill slits. Mouth large and parabolic, ventral on head; jaws moderately protrusable but notgreatly distensible laterally. Teeth large, anteriors and laterals narrow and awl-shaped or blade-like to broad, compressedand triangular, in 22 to 31/20 to 29 (43 to 60 total) rows; two rows of large anterior teeth on each side in upper jaw, separatedfrom the smaller upper lateral teeth by one row of small intermediate teeth on each side; three rows of lower anterior teeth oneach side, the first two variably enlarged but the third about as large as laterals; no symphysial teeth. Trunk fusiform andmoderately slender to very stout, firm and not flabby. Caudal peduncle strongly depressed and with strong, high keels andboth upper and lower crescentic precaudal pits. Dermal denticles very small and smooth, with flat crowns, small ridges andcusps and with cusps directed posteriorly on lateral denticles. Pectoral fins very long and narrow, shorter to somewhatlonger than head in adults; pectoral-fin skeletons plesodic with radials extending far into fin webs. Pelvic fins small, muchsmaller than first dorsal fin but larger than second dorsal and anal fins; pelvic-fin skeleton aplesodic, not extending into finweb. First dorsal fin large, high, erect and angular or somewhat rounded; fin skeleton semiplesodic, extending partway into

Sharks of the World, Vol. 2 97fin web. Second dorsal and anal fins minute, much smaller than first dorsal fin, with narrow pivoting bases. Caudal fin lunate,dorsal lobe moderately long, less than one third as long as rest of shark, ventral lobe long and strong, nearly as long as upperlobe. Neurocranium moderately high, not compressed, with moderately long rostrum, depressed internasal septum andwidespread nasal capsules, large orbits with strong supraorbital crests, greatly enlarged stapedial fenestrae, and posteriorlyexpanded hyomandibular facets. Vertebral centra strongly calcified, with well-developed double cones and radii but withoutprominent annuli. Total vertebral count 153 to 197, precaudal count 85 to 114, diplospondylous caudal count 66 to 86.Intestinal valve of ring type with 38 to 55 turns. Size moderately large to very large with adults 1.8 to about 6 m long.Distribution: Lamnids have a broad geographic distribution in virtually all seas except where the ice pack covers the Arcticand Antarctic Oceans. One of them, the white shark (Carcharodon carcharias) has one of the widest ranges of anycartilaginous fish. The salmon shark (Lamna ditropis) has the most limited distribution in the family, the temperate andboreal North Pacific and Arctic fringe, but lives in an enormous area.Habitat: Lamnids are tropical to boreal or notal, littoral to epipelagic sharks in continental and insular waters from the surfline and intertidal of shallow bays to the outer shelves and open ocean and rarely down the slopes to at least 1 280 m.Lamnids are apparently intolerant of fresh water and do not ascend rivers. Salmon sharks and porbeagles (Lamna) aretolerant of cold and moderate water temperatures but avoid the tropics, while makos (Isurus) are restricted to tropical andtemperate seas. Carcharodon broadly overlaps the habitats of Lamna and Isurus.Biology: The mackerel sharks are fast-swimming, active pelagic and epibenthic swimmers, some of which are capable ofswift dashes and spectacular jumps when chasing their prey. Mackerel sharks are partially warm-blooded, and have amodified circulatory system that enables them to retain a body temperature warmer than the surrounding water. This permitsa higher level of activity and may increase the power of their muscles as well as allow some of them to be active in coldtemperatures. They feed on a wide variety of bony fishes, other sharks, rays, chimaeras, marine birds and reptiles, seals andsea lions, whales and dolphins, squid, bottom crustaceans and molluscs, carrion, and occasionally terrestrial vertebrates(rarely including humans). Development is ovoviviparous, without a yolk-sac placenta. Like other lamnoids these sharkshave uterine cannibalism, in which developing foetuses feed on fertilized eggs (oophagy) for most of their gestation period.More than one foetus survives in each uterus in most species, and it is not known if intrauterine siblings will attack anddevour one another as in Carcharias taurus. All the living species of lamnids are of large size, with a maximum length of 3.0to perhaps 6mormore. A giant, rather recently extinct (late Pliocene) member of the white shark genus (the megatoothshark, Carcharodon megalodon, often placed in other genera or even families by some modern palaeontologists) attainedan estimated length when adult of about 11 to 20 m (Gottfried, Compagno and Bowman, 1996). It was one of the largest andmost powerful predatory vertebrates ever to live, rivalled only by certain cetaceans (sperm whales, orcas and giantarchaeocetes), the largest pliosaurs and mosasaurs, giant Mesozoic and Cenozoic crocodiles, and the largest terrestrialtheropod dinosaurs.Interest to Fisheries and Human Impact: These sharks are important objects of oceanic and offshore continental fisheriesbecause of their fine meat, but are also utilized for their oil, fins, hides, fishmeal, jaws and teeth. Some species are oceanic inwhole or part, and are mainly taken with pelagic longlines and gill nets. Anchored gill nets and trammel nets, fish traps,hook-and-line, harpoons, and pelagic and bottom trawls can capture these sharks.Porbeagles and salmon sharks (Lamna) are apparently inoffensive and have not bitten people in the water. The shortfinmako (Isurus oxyrinchus) has bitten swimmers and divers on a few occasions but may often display and stagemock-charges while confronted by divers. More often shortfin makos have bitten boats, especially after being provoked byhooking. The longfin mako has never been known to bite people or boats. However, this family contains what is generallyconsidered the most ‘dangerous’ shark, the white shark (Carcharodon carcharias), because of its regular if infrequent habitof biting swimmers, divers, surfers and boats.Local Names: Mackerel sharks, Man-eater sharks, Man-eating sharks, Man-eaters, Great white sharks, Mako-sharks,Porbeagles (English); Lamie, Requin-bleu, Taupe, (France); Heringshai, T’u sha k’o; Akuly sel devye or Sel devye akuly(Russia); Nezumizame-ka (Japan); Anequins (Mozambique).Remarks: This account follows Compagno (1984, 1990b, 1999) in recognizing a single family for the genera Carcharodon,Isurus and Lamna, which is termed Lamnidae or Isuridae by most authors. Lamnidae has priority, stemming from Müllerand Henle’s (1838a, 1839) family Lamnoidea or Lamnae, and being recognized by numerous authors including Bonaparte(1838, 1839, subfamily Lamnini), Müller (1845), Bleeker (1859), Gill (1862b, 1872), Owen (1866), Günther (1870), Hasse(1879), Jordan and Gilbert (1883), Woodward (1889), Jordan and Evermann (1896), Regan (1906a), Goodrich (1909),Bridge (1910), Engelhardt (1913), Jordan (1923), Bertin (1939a), Berg (1940), Fowler (1947, 1967a), Berg and Svetovidov(1955), Matsubara (1955), Patterson (1967), Blot (1969), Bailey et al. (1970), Lindberg (1971), Pinchuk (1972), Compagno(1973, 1981b, 1982, 1984, 1999), Nelson (1976, 1984, 1994), Applegate et al. (1979), Gubanov, Kondyurin and Myagkov(1986), Cappetta (1987), Carroll (1988), Eschmeyer (1990, 1998), Robins et al. (1991a), and Helfman, Collette and Facey(1997). Isuridae was apparently not used until Gill (1893) essentially revived Gray’s (1851) Isurini as a family. Although anunjustified replacement of Lamnidae the family Isuridae was used by a number of influential authors, including Garman(1913), Lozano y Rey (1928), White (1936, 1937), Whitley (1940), Fowler (1941), Romer (1945, 1963), Bigelow andSchroeder (1948), Schultz and Stern (1948), Smith (1949), Arambourg and Bertin (1958), Garrick and Schultz (1963),Budker and Whitehead (1971), Bass, D’Aubrey and Kistnasamy (1975a), and Chu and Meng (1979).

98 FAO Species Catalogue for Fishery Purposes No. 1Whitley (1940) separated the white shark (Carcharodon) in its own family Carcharodontidae as distinct from Isuridae(Isurus and Lamna). This was independently proposed by the innovative Soviet palaeontologist L.S. Glikman (1964, 1967),who also placed Lamna in a subfamily Lamninae of the family Odontaspididae. In addition, Glikman separated living fossilspecies of Isurus into two genera, Isurus and Lamiostoma, and placed these respectively in the families Isuridae andLamiostomatidae. Glikman’s arrangement was followed by Rass and Lindberg (1971), while Lindberg (1971) recognizedLamiostomatidae as distinct from Lamnidae. Glikman (1964) suggested that the living lamnid genera were independentlyderived from the basal family Odontaspididae, and so required separate families. However, phyletic studies utilizingmorphology (Maisey, 1985; Compagno, 1990b; Long and Waggoner, 1996) and DNA sequencing (Martin and Naylor, 1997;Naylor et al., 1997) suggest that the Lamnidae is a monophyletic group for the living Carcharodon, Isurus and Lamna,which is followed here.Günther (1870), Regan (1906a) and Engelhardt (1913) included Isurus or its synonym Oxyrhina as a synonym of Lamnawhile Garman (1913), White (1936, 1937), and Fowler (1941) synonymized Lamna with Isurus. Most modern authors followMüller and Henle (1839) and more recently Bigelow and Schroeder (1948) in considering Isurus and Lamna as separategenera, which is supported by external morphology, dentition and anatomical studies (Compagno, 1990b; Long andWaggoner, 1996). Gray (1851) recognized both genera but used Isurus as a synonym of Lamna and Oxyrhina in place ofIsurus.Many earlier authors included the basking shark (Cetorhinus) and the threshers (Alopias) in the family Lamnidae. Modernauthors generally recognize Lamnidae (or Isuridae), Cetorhinidae and Alopiidae as distinct families, although Shirai (1996)recently demoted the Lamnidae as a subfamily Lamninae of the Alopiidae along with the subfamilies Cetorhininae andAlopiinae.Literature: Garman (1913); Fowler (1941, 1967a); Bigelow and Schroeder (1948); Garrick and Schultz (1963); Farquhar(1963); Shiino (1972, 1976); Bass, D’Aubrey and Kistnasamy (1975a); Compagno (1984, 1990b); Alexander (1998).Key to Genera:1a. Teeth serrated, uppers flat and withbroadly triangular cusps (Fig. 68) . . . . Carcharodon1b. Teeth smooth-edged, uppers notgreatly flattened and with narrowlytriangular cusps (Fig. 69a) . . . . . . . . . . . . 2serratedUPPER AND LOWER TOOTHFig. 68 Carcharodon2a. Lateral cusplets present on mostteeth (sometimes absent in young)(Fig. 69a); origin of first dorsal fin oversmoothor anterior to inner margins of pectoralfins; origin of second dorsal fin overorigin of anal fin; a secondary keelpresent below main keel on caudal fin(Fig. 69b) . . . . . . . . . . . . . . . . . . . Lamna2b. No cusplets on teeth (Fig. 70a); originof first dorsal fin over or behind reartips of pectoral fins; origin of seconddorsal fin well in front of anal-fin origin;no secondary keel on caudal fin(Fig. 70b) . . . . . . . . . . . . . . . . . . . Isuruscuspletsa) UPPER TOOTHa) UPPER TOOTHb) LATERAL VIEWFig. 69 Lamnab) LATERAL VIEWFig. 70 IsurusCarcharodon Smith, 1838Genus: Carcharodon Smith, in Müller and Henle, 1838a, Mag. Nat. Hist., new ser., 2: 37. Placed on the Official List ofGeneric Names in Zoology (Name no. 1658) by the International Commission on Zoological Nomenclature (1965, Opinion723.3b, Bull. Zool. Nomencl., 22(1): 32).

Sharks of the World, Vol. 2 99Type Species: Squalus carcharias Linnaeus, 1758, by subsequent monotypy through Carcharias lamia Rafinesque, 1810(International Commission on Zoological Nomenclature, 1965, loc. cit.).Number of Recognized Species: 1.Synonyms: Subgenus Carcharias Cuvier, 1816 (Genus Squalus Linnaeus, 1758): 125, in part. Placed on the List ofRejected and Invalid Generic Names in Zoology (Name no. 811) by the International Commission on ZoologicalNomenclature (1965, Opinion 723.5c: 33). Cuvier’s Carcharias had only three species, Squalus carcharias, S. vulpes(= Alopias vulpinus), and S. glaucus (= Prionace glauca). S. carcharias is the type of Carcharias Cuvier by absolutetautonymy, but is a junior homonym of Carcharias Rafinesque, 1810 (Carcharias taurus, type species by monotypy).Carcharias Cuvier was used extensively for carcharhinids following Müller and Henle (1839: 27). Genus CarcharodenBleeker, 1860: 58, in combination Carcharoden rondeletii. Apparent misspelling for Carcharodon, as it is spelled correctlyin that paper on p. 57 (Carcharodon capensis).Diagnostic Features: Snout bluntly conical. Eyes small, 0.7 to 1.8% of total length. Nostrils situated adjacent to head rim inventral view. Mouth width 1.1 to 2.3 times its length. Anterior teeth enlarged; anterior, intermediate and lateral teethcompressed and forming a continuous cutting edge; intermediate teeth enlarged and over two-thirds height of adjacentanteriors, with reversed cusps that are directed anteromesially; second lower anterior teeth moderately enlarged and aboutas high or usually lower than second upper anterior tooth; total tooth count 44 to 52; roots of anterior teeth broadly arched,with root lobes broad and not elongated; lateral cusplets present only on teeth of smaller sharks below about 3 m long butlost in adults; teeth with serrated edges; cusps of anterior teeth not strongly flexed. Body usually stout. First dorsal-fin originusually over the pectoral inner margins. Anal-fin origin under or slightly posterior to second dorsal-fin insertion. Secondarycaudal keels absent. Total vertebral count 170 to 187. Cranium with rostral cartilages not swollen and hypercalcified.Intestinal valve count 47 to 55. Length of adults 3.8 to almost 6 m and possibly longer. Usually a black axillary spot at pectoralfin insertions; pectoral fin tips usually abruptly black on their ventral surfaces.Remarks: The white shark (Carcharodon carcharias) is apparently the sole living species of this genus although manyfossil species are also recognized. The white shark was often confused with requiem sharks (Carcharhinidae) and othersharks by eighteenth and early nineteenth century writers, and was often placed in the genus Carcharias along with otherlarge lamnoid and carcharhinoid sharks. Smith (1838a) proposed Carcharodon as a unique genus for the white shark, butwithout allocating any species. Müller and Henle (1839) recognized a single species, Carcharodon rondeletii Müller andHenle, 1839. The genus was accorded essentially universal recognition by subsequent authors, but the type species wasquestionable and was eventually stabilized by a ruling of the International Commission on Zoological Nomenclature (1965).Bigelow and Schroeder (1948), White, Tucker and Marshall (1961), and the International Commission on ZoologicalNomenclature (1965) give further details of the complex nomenclatural history requiring this ruling.There have been a few attempts to name regional species of white sharks and distinguish separate regional populations.Smith (1849) proposed a new species, C. capensis, from South Africa, while Whitley (1939) separated Australian whitesharks as C. albimors. Neither of these were especially characterized, and are generally synonymized with C. carcharias.More recently Bass, D’Aubrey and Kistnasamy (1975a) distinguished South African white sharks from those from theNorthern Hemisphere by coloration and vertebral count. These authors noted that the axillary spots reported on westernAtlantic white sharks (Garman, 1913; Bigelow and Schroeder, 1948) were absent in specimens they examined. At least oneof the specimens examined by Bass, D’Aubrey, and Kistnasamy (1975a, fig. 10) lacked axillary spots, but Smith (1849),Smith (1951), and D’Aubrey (1964) illustrated South African white sharks with axillary spots. The writer has examined manywhite sharks specimens with axillary spots from South Africa and California, United States, while axillary spots were presenton several live white sharks filmed at Dyer Island and Struis Bay, South Africa. Axillary spots may be absent in white sharksfrom Argentina (Siccardi, Gosztonyi and Menni, 1981) and were poorly defined on one examined by the writer fromCalifornia. Apparently the absence of axillary spots cannot be used to distinguish South African white sharks.Bass, D’Aubrey and Kistnasamy (1975a) suggested that lower counts of precaudal vertebral centra (100 to 108, mean= 103.2, n = 24) might distinguish white sharks from Natal as a separate population from California white sharks (103 to 108,mean = 105.2, n = 14; California data from Springer and Garrick, 1964). Further comparison of precaudal vertebrae countsfor Californian and South African specimens by the author (Compagno unpub. data; n = 56) verified a small but statisticallysignificant difference in means between the samples, but the small sample sizes, broad overlap in ranges and standarddeviations in the vertebral count samples, do not rule out the differences as being a result of sampling error. Siccardi,Gosztonyi and Menni (1981) presented precaudal counts for two Argentinean white sharks (a female with 104 centra and amale with 105), which fall within the range of the South African and Californian samples.It is not obvious from available data on morphometry, meristics, coloration and skeletal anatomy that white sharks fromdifferent ‘centres of abundance’ are recognizably separable. The wide distribution and habitat of the white shark suggests asingle species, but discrete centres of abundance and concentration of breeding areas in warm-temperate coastal seassuggest discrete populations or subpopulations with potential genetic interchange via wide-ranging adults. There areseveral projects underway to compare DNA of white sharks from various areas to determine possible populationaldifferences.

100 FAO Species Catalogue for Fishery Purposes No. 1Carcharodon carcharias (Linnaeus, 1758) Fig. 71Squalus carcharias Linnaeus, 1758, Syst. Nat., ed. 10, 1: 235. Placed on the Official List of Specific Names in Zoology (Name no.2056) by the International Commission on Zoological Nomenclature, (1965, Opinion 723.4b, Bull. Zool. Nomencl., 22(1): 32).Holotype unknown, type locality “Europa”. Also no types known according to Eschmeyer (1998, Cat. Fish.: CD-ROM).Synonyms: Carcharias lamia Rafinesque, 1810b: 44. Type locality: Sicily. No types? Placed on the List of Rejected and InvalidSpecies Names in Zoology (Name no. 811) by the International Commission on Zoological Nomenclature, (1965, Opinion 723.6:33). ?Squalus (Carcharhinus) lamia Blainville, 1816: 121 (in part?). Name only. Carcharias verus Cloquet, 1817: 69. Europe. Notypes? ?Squalus (Carcharhinus) lamia Blainville, 1825: 88, pl. 22, fig. 2 (in part?). No types? Carcharias rondeletti Bory deSaint-Vincent, 1829: 596. Europe. Types? Squalus (Carcharias) vulgaris Richardson, 1836: 288. All seas. An unexplained newname, without types, according to Eschmeyer (1998: CD-ROM). Carcharodon smithii Müller and Henle, in Agassiz, 1838: 91.Name only, attributed to teeth from two jaws in pl. F, fig. 3; dentition pictured shows some overlap and no intermediate, quite possiblya Carcharhinus. Type locality: Possibly South Africa, presumably named after Andrew Smith and perhaps a nomen nudum.Carcharodon smithii was not mentioned in Müller and Henle (1838a, 1839, 1841) though these authors (1839: 70) note dentitionaldifferences between a specimen collected by A. Smith and other material and suggest that there may be two species. Carcharodonsmithii Müller and Henle, 1839: 9 (in synopsis, name only). Type locality: Probably South Africa, and presumably named afterAndrew Smith. Apparently a nomen nudum. Note difference in spelling from Agassiz, 1838: 91. Carcharodon rondeletii Müller andHenle, 1839: 70. Type locality: “Mittelmeer, atlantischer Ocean, Kap, stilles Meer.”. This appears to be an original description and notmerely a reallocation of Carcharias rondeletti Bory de Saint-Vincent, 1829, to Carcharodon. Syntypes: A stuffed specimen(holotype of Carcharodon capensis) belonging to Andrew Smith and now in the British Museum (Natural History), a specimen in theSenkenberg Museum from the Adriatic Sea, a jaw in the Zoologisches Museum, Museum für Naturkunde der Humboldt-Universität,Berlin, and two specimens in the “United Services Museum” (US National Museum of Natural History, Eschmeyer, 1998: CD-ROM),from the Pacific Ocean. Carcharias atwoodi Storer, 1848: 72. Type locality, Provincetown, Massachusetts. Holotype: Museum ofComparative Zoology, Harvard, MCZ 89505 (missing), jaws may be MCZ 775-S (Eschmeyer, 1998: CD-ROM). Carcharodoncapensis Smith, 1849: pl. 4 and accompanying text, pages not numbered. Holotype and only specimen: 2.135 m (7 ft) TL stuffedfemale, from Cape Seas, South Africa, deposited in the Museum of the Zoological Society of London, but according to Gray (1851:61), subsequently donated to the British Museum (Natural History). The holotype, BMNH 1850.9.5:3, is still extant and in goodcondition and was located and examined on 10 June 1994 in the BMNH collections; current TOT (extended TL) is about 2 035 mm.Carcharias vorax Owen, 1853: 94. Holotype? No locality, identity uncertain, based on vertebrae and teeth of a 7.65 m (23 ft)specimen. Carcharias maso Morris, 1898: 412. Australia. Possible nomen nudum according to Eschmeyer (1998: CD-ROM). NotSqualus (Carcharias) maou Lesson, 1830 = Carcharhinus longimanus (Poey, 1861). Carcharodon albimors Whitley, 1939: 240.Holotype: Australian Museum, Sydney, AMS I.1723 (Paxton et al., 1989: 67; Eschmeyer, 1998: CD-ROM), New South Wales,Australia.Other Combinations: None.FAO Names: En - Great white shark; Fr - Grand requin blanc; Sp - Jaquetón blanco.UPPER AND LOWER TEETH ON LEFT SIDEFig. 71 Carcharodon carchariasUNDERSIDE OF HEAD

Sharks of the World, Vol. 2 101Field Marks: Heavy spindle-shaped body, moderately long conical snout, huge, flat, triangular, serrated blade-like teeth,long gill slits, large first dorsal fin with dark free rear tip, minute, pivoting second dorsal and anal fins, strong keels on caudalpeduncle, no secondary keels on caudal base, crescentic caudal fin. Colour: dorsal surface lead grey or brownish grey toblackish above, ventral surface of body white, iris of eye conspicuously black, margin between dorsal dark and ventral whitesurfaces sharply delimited.Diagnostic Features: See genus Carcharodon above.Distribution: Wide-ranging in most seas. Western Atlantic: Newfoundland to Florida, Bahamas, Bermuda, Cuba, northernGulf of Mexico; also Brazil and Argentina. Eastern Atlantic: Possibly England, also France and Bay of Biscay, to Gibraltar,the entire Mediterranean Sea (absent from Black Sea), Madeira, Canary Islands, Senegal, Gambia, Ghana, possibly Zaire,Angola, Namibia, South Africa (Northern and Western Cape Provinces); also Gough Island. Indo-West Pacific: South Africa(Eastern Cape and KwaZulu-Natal Provinces), Mozambique, Tanzania (Zanzibar), Kenya, Seychelles, Madagascar,Mauritius, possibly Red Sea and Persian Gulf (Kuwait?), Sri Lanka, possibly Indonesia, Australia (Queensland, New SouthWales, Victoria, Tasmania, South and Western Australia), New Zealand (including Norfolk, Stewart, and Chatham Islands),New Caledonia, Philippines (Mindanao, Palawan), China, Taiwan (Province of China), Japan, North Korea, South Korea,Russia (Siberia, possibly Sea of Okhotsk and Bering Sea), Bonin Islands (Tanna Island). Central Pacific: Marshall Islands,Hawaiian Islands, open ocean between Polynesia and South America. Eastern Pacific: Bering Sea and Gulf of Alaska toGulf of California, including Canada (British Columbia) and the entire Pacific coast of the USA (Washington, Oregon,California, Alaska), and much of Mexico, also Panama, Ecuador, Peru, Chile, and Galapagos Islands.Habitat: This huge and formidable shark is best known as a coastal and offshore inhabitant of the continental and insularshelves in temperate seas, but it readily penetrates the epipelagic zone and occurs in the inshore equatorial tropics and athigh latitudes. The presence of large individuals off oceanic islands far from land and where breeding does not occur (e.g.,Hawaiian Islands, Gough Island, Mauritius, Seychelles) as well as recent open-ocean gill net records in the North Pacificsuggest that it regularly occurs in the epipelagic zone although less commonly than smaller, more abundant species ofIsurus and Lamna. It is seldom recorded from pelagic longline catches unlike other lamnids. This may be a function ofrelative rarity in the epipelagic zone and gear selectivity, with larger animals breaking off gangions and the species seldombeing caught or reported in the past.The white shark often occurs close inshore to the surfline and even penetrates shallow bays, estuaries and the intertidalzone in continental coastal waters, but also frequents offshore continental and oceanic islands (especially those withpinniped colonies) and inshore and offshore fish banks. The white shark can be found at the surface down to the bottom inepicontinental waters but rarely ranges down the continental slope, where it was once caught on a bottom longline at1 280 m along with the large sixgill shark (Hexanchus griseus). More recently it has been taken in bottom trawls on the outershelf down to 130 m off South Africa. It is often seen by divers off rocky reefs and near shipwrecks but in the tropics it isoccasionally sighted on coral reefs. The white shark has one of the widest habitat and geographic ranges of any fish-likevertebrate, and readily tolerates temperature extremes from the Bering Sea and sub-Antarctic islands to the inshore tropics.It apparently does not occur in fresh water, but can be expected to occur in most marine environments if only sporadically.Biology: This species is a very active, nomadic, social shark with a fluid, powerful, scombroid-like mode of cruising thatallows it to efficiently cruise and manoeuvre for long periods at a relatively slow speed. Tracking of white sharks off the east

102 FAO Species Catalogue for Fishery Purposes No. 1coast of the United States and south Australia with sonic tags indicated an average cruising speed of about 3.2 kph, with oneshark covering 190 km in 2.5 days. The white shark is capable of sudden high-speed dashes and drastic manoeuvres. Itsometimes jumps (breaches) right out of the water in pursuit of prey, and jumps to attack prey at the surface, in midair, oreven on rocks above the water. It can shoot straight out of the water like a dolphin for no obvious reason.The white shark is most commonly reported from cold and warm temperate ‘centres of abundance’, though there are enoughtropical continental and oceanic records to suggest that at least larger individuals have a wide temperature range and readilypenetrate the tropics. Smaller individuals, below 3 m long, may be mostly restricted to temperate continental seas, and thedistribution of presumably newborn individuals in the 100 to 160 cm size range suggest that pupping and nursery grounds forthe species are also in temperate to subtropical coastal and mostly continental waters. Known centres of abundance,including nursery and probably breeding areas, include the west coast of the United States (California), Mexico (BajaCalifornia, possibly Gulf of California), the Mid-Atlantic Bight off the USA (New England and Mid-Atlantic States), theMediterranean Sea, the east coast of South Africa (from False Bay to KwaZulu-Natal), southern Australia, New Zealand,and Japan.White sharks are endothermic and by development of countercurrent vascular heat exchangers maintain highertemperatures in their body musculature, brains, eyes and viscera than the surrounding water (Carey and Teal, 1969; Careyet al., 1982, 1985; Carey, 1982, 1990; Tricas and McCosker, 1984; Block and Carey, 1985; McCosker, 1987; Goldman et al.,1996; Goldman, 1997). Body muscle may run 3 to 5°C higher than ambient (with few sharks measured and possibleproblems with measuring gear), but stomach temperatures can be 10 to 14°C above ambient and relatively constant in coldwater. The heat retention system of the white shark (and possibly its large size) may be particularly advantageous in allowingit to function efficiently as a fast, agile predator in cold water, and to hunt active large prey.Relatively little is known of the abundance of this species, except that it is uncommon to rare compared to most other sharkswhere it lives, even in temperate coastal waters. Catches in some areas have been as many as 50 to 100 per year (SouthAfrica and South Australia) in the past but mostly less in others. There have been unsubstantiated claims that the species isincreasing in numbers in some areas (off central California and South Africa), as a result of increasing numbers of pinnipeds.There is no hard evidence to prove this, and increasing fishing pressure and injuries from targeted and bycatch fisheries insuch areas may be very well having the reverse effect. Declines in shark-meshing catches of white sharks have occurred offQueensland and New South Wales, and reductions in sightings of white sharks in Spencer Gulf, South Australia may havebeen related to commercial and sports fishing mortality.Pronounced periodicity in white shark abundance may occur in some areas, apparently correlated with temperature and tosome extent with life stage, or by movements of individuals or groups in response to prey concentrations or other stimuli.Also, shifts may occur in size and gender composition of white sharks off ‘white shark sites’ such as fish banks or sealcolonies where the sharks congregate. In colder, higher latitudes at the periphery of its range in North America, the whiteshark moves into more northern areas when water masses warm up in the summertime. Off KwaZulu-Natal, South Africa, itwas believed that individuals below 2.8 m long segregated themselves from larger individuals and moved out of the areawhen temperatures rose above 22°C, but this was proven wrong by more extensive data presented by Cliff, Dudley andDavis (1989). Essentially both size classes are present all year round in the area, although numbers of both fluctuated bytime of year and area. In central California (Monterey Bay) white sharks are present year round but are slightly commonerwhen water temperatures rise to 14 to 15°C than when it is below 11°C.Observation of white sharks at ‘white shark sites’ suggest that they are nomadic and may spend relatively short periods ofless than a day at a given site, but at least some individuals revisit these sites periodically, from a few days to over severalyears. Recognizable individuals have been photographed and resighted for several years off the Farallon Islands,California. Streamer and plate tagging off Australia and South Africa, and sonic tagging off Australia also revealed short tomoderately long-term site specificity of some individuals. Tagging of white sharks and photographing of dorsal fins and bodypatterns (‘bodyprinting’) have shown fairly long-range movements, including between northern and southern California(over 700 km, Anderson and Goldman, 1996) and between Dyer Island and Mossel Bay (about 300 km) or from Dyer Islandto KwaZulu-Natal, South Africa (over 1 100 km, M. Marks and L.J.V. Compagno, unpub. data). So far there have been nointercontinental resightings or recaptures of tagged or photographed sharks, but given the size and strong swimmingabilities of this shark and scattered records in the tropics and off oceanic islands these are to be expected and may berevealed by usage of improved tracking technologies and by development of ‘bodyprint’ libraries that could be made widelyavailable via the Internet.The white shark occurs singly or in pairs but is apparently a social animal that can be found in aggregations or congregationsof 10 or more; polarized schooling possibly does not occur (apart from parallel swimming of two individuals) but sharks in agroup may mill about and interact socially while showing various behaviours, often one-on-one but sometimes with moreindividuals involved. White sharks are ‘inquisitive’ animals, and often closely and repeatedly investigate human activities(including divers). Behaviour and sociobiology of this species is sketchily known at present but it is apparently at least ascomplex as the better-known bonnethead shark (Sphyrna tiburo), with most of the noncourtship, social and asocialbehaviours reported by Myrberg and Gruber (1974) being present in the white shark along with several additionalbehaviours not observed in the bonnethead. A detailed account of these behaviours will be published elsewhere. A possiblyagonistic behaviour commonly directed at people underwater or above the surface is gape, a graded distension andprotrusion of jaws also seen in the shortfin mako (Strong, 1996). Spy-hopping, in which the shark raises its head out of thewater (often alongside a boat), and tail-slap, in which the shark raises its tail and caudal fin out of the water and smashes itagainst the surface, are common behaviours. Klimley, Pyle and Anderson (1996) suggest the latter is a social signal

Sharks of the World, Vol. 2 103between contesting sharks over a prey item, which is plausible from their evidence although tail-slapping also occurs insituations of ambiguity as when a shark has a bait pulled out of its grasp and may also be directed toward inanimate objectsor even people. White sharks will hunch, arching their backs and depressing their pectoral fins in a possibly agonistic displayas in bonnetheads and grey reef sharks (Carcharhinus amblyrhynchos). One white shark may closely follow another, andtwo approaching white sharks on a collision course may give-way, with one shark avoiding the other, or both may give-wayalmost simultaneously (stand-back). Follow give-way is sometimes seen at baits, where a shark approaching a bait isfollowed by a second (sometimes larger); the first shark suddenly aborts its approach and the second eats the bait, or a thirdshark may follow and displaces the second. Courtship behaviour is poorly known in the white shark but rows of limited toothmarks on the pectoral fins of large adult female sharks suggests that complex courtship behaviour may be present as in thesand tiger shark (Carcharias taurus; Gordon, 1993).Territoriality in the white shark (as contrasted with site specificity) cannot be demonstrated at present, but there is someevidence for sorting of individuals into a partly size-related hierarchy around food sources such as dead whales, pinnipedcolonies or feeding stations provided by people. White shark tooth scratches and inhibited bite marks are seen onindividuals of all sizes, both sexes, and all postnatal maturity stages. They have been interpreted as evidence of intraspecificconflict, possibly in competition for food resources, but they probably have a broader context of social interactions. In certainareas (southern Australia, the south coast of South Africa, and central California), white sharks may have habituated tohuman-provided food sources and may have learned to seek out fishing boats to exploit hooked fish or the baits provided byshark cage-dive operators and film-makers.The white shark is ovoviviparous (aplacental viviparous) and practices uterine cannibalism as do many other lamnoids,apparently in the form of embryos and foetuses eating large numbers of nutritive eggs (oophagy). The gestation period is notknown at present but could be a year or more, and may or may not include taking a year off to recuperate. There are fewrecords of pregnant females and litter size, which varies from 2 to possibly 14 young. Females mature between 4 and 5 mlong and 12 to 14 years old and reach at least 23 years old, while males mature between 3.5 and 4.1 m long and 9 or 10years old (ageing from growth rings on vertebral centra, assumed to be annual); the maximum age of either sex is unknownbut has been roughly estimated at about 27 years from a von Bertalanffy growth model (Cailliet et al., 1985) assuming amaximum size of 7.6 m and indeterminate growth to that size.The rarity of pregnant female white sharks could be explained in part by spatial separation from other white sharks duringpregnancy (without evidence, however) and their sheer size that precludes capture by most fishing gear. However, it ispossible that adult female white sharks are uncommon (more so than adult males), and possible also that they may havevery low fecundity, with only a few adult females being pregnant at any one time and with an interval of a few years betweenpregnancies.The white shark readily scavenges on available carrion, garbage, and fish caught on lines. This species probably obtainsmost of its prey by killing it, but is highly opportunistic as with terrestrial apex predators. Its diet in different areas may varyaccording to the availability (including abundance) and vulnerability of suitable prey, as well as by motivation of the predator.Live prey of individual white sharks is mostly marine vertebrates and invertebrates smaller than itself, ranging in size fromsmall schooling fishes and squid to elephant seals and grey whale calves. The chief prey categories of the white shark areray-finned bony fishes (Actinopterygii), cartilaginous fishes, marine mammals, marine birds, cephalopods, and crustaceans,with marine reptiles (sea turtles) and gastropods being relatively unimportant.Bony fish prey of the white shark includes a wide range of large and small, demersal and pelagic forms, such as sturgeon(Acipenseridae), menhaden and pilchards (Clupeidae), salmon (Salmonidae), sea catfish (Ariidae), lings (Gadidae), hake(Merluccidae), flounders and halibut (Paralichthyidae), rockfish (Sebastes spp., Scorpaenidae), cabezon (Scorpaenichthysmarmoratus, Cottidae), lingcod (Ophiodon elongatus, Hexagrammidae), barracuda (Sphyraenidae), striped bass(Morone saxatilis, Percichthyidae), bluefish (Pomatomidae), butterfish (Stromateidae), grunters (Pomadasyidae),croakers (Sciaenidae), garrick and maasbanker (Lichia and Trachurus, Carangidae), porgies or sea bream (Sparidae),mackerels and tuna including bluefin tuna (Thunnus thunnus, Scombridae), swordfish (Xiphias gladius, Xiphiidae), andocean sunfish (Mola mola, Molidae). White sharks are known to congregate at concentrations of schooling bony fishessuch as pilchards and bluefish, and follow the KwaZulu-Natal sardine (Sardinops) run off South Africa.Chondrichthyan prey of the white shark includes other sharks such as shortfin mako (Isurus oxyrinchus, Lamnidae), sandtiger sharks (Carcharias taurus, Odontaspididae), catsharks (Scyliorhinidae), houndsharks (Galeorhinus, Mustelus,Triakidae), requiem sharks (Carcharhinus, Prionace, Rhizoprionodon, Carcharhinidae), hammerheads (Sphyrna,Sphyrnidae), and spiny dogfish (Squalus, Squalidae). Basking shark (Cetorhinus) meat has been found in several whitesharks, apparently taken as carrion from harpooned sharks. Whale shark (Rhincodon) remains were found in one sharktaken in a beach meshing net. It is presently unknown if the white shark ever attacks adult basking or whale sharks thoughsmaller juveniles of both might be readily killed and eaten. Batoid prey includes giant guitarfish (Rhynchobatus,Rhynchobatidae), guitarfish (Rhinobatos, Rhinobatidae), skate egg cases (Rajidae), stingrays (Dasyatidae), and eagle rays(Myliobatis and Pteromylaeus, Myliobatidae). Chimaeras (Chimaeriformes) are eaten and include shortnose chimaeras(Chimaeridae), and elephantfish (Callorhinchidae). No small white sharks have been found in white shark stomachs,although large white sharks will eat shortfin makos and other very fast prey, will attack hooked, injured conspecifics, and willdeliver inhibited bites to other white sharks. The writer suspects that white sharks have behavioural inhibitions oncannibalization under ordinary circumstances, as reflected by known prey records and from social interactions of small andlarge white sharks within aggregations.

104 FAO Species Catalogue for Fishery Purposes No. 1Sea turtles are occasionally eaten by the white shark, including loggerhead and green turtles (Carettidae) and leatherbackturtles (Dermochelyidae) but apparently not to the degree that the tiger shark (Galeocerdo) preys on them (Fergusson,Marks and Compagno, 2000).Marine birds fall prey to white sharks and include cormorants (Phalacrocoracidae), gannets (Sulidae), gulls (Laridae), giantpetrels (Macronectes, Procellariidae), pelicans (Pelicanidae), and penguins (Spheniscidae). Jackass penguins(Spheniscus demersus) are commonly grabbed, cut or slashed by white sharks off South Africa and are often killed butseldom eaten. White sharks also use their snouts to flip or bounce seabirds on the surface without eating them, and victimshave included gulls (Laridae) and shearwaters and fulmars (Procellariidae).Marine mammals can be an important food source for white sharks in some areas, and these include a number of cetaceansand pinnipeds. Those cetaceans killed and eaten include harbor porpoises (Phocaena phocaena, Phocaenidae), commondolphins (Delphinus delphis) and bottlenose dolphins (Tursiops sp., Delphinidae), and even a grey whale calf (Estrichtiusrobustus). White shark inhibited bite marks have been seen on striped dolphins (Stenella caerulalba) from South Africa andon a pygmy sperm whale from California (Long, 1991). White shark predation on odontocetes is more difficult to study thanpinniped predation, though careful study of stranded cetaceans can reveal greater diversity of cetacean predation,scavenging, and non-predatory biting by white sharks. Dead baleen whales and other large cetaceans may contribute asignificant amount to the white shark’s diet in some areas (Long and Jones, 1996), but such food is sporadically available atbest.True, earless or ‘hair’ seals (Phocidae) taken by white sharks include harbor seals (Phoca vitulina), grey seals(Halichoerus grypus) and northern elephant seals (Mirounga angustirostris), with predation suspected for leopard seals(Hydrurga leptonyx), Hawaiian and Mediterranean monk seals (Monachus), and southern elephant seals (Miroungaleonina). Eared seals (Otariidae) taken include Steller’s sea lion (Eumetopias jubata), California sea lions (Zalophuscalifornianus), South African fur seals (Arctocephalus pusillus pusillus), Australian fur seals (A. p. doriferus), SouthAmerican fur seals (A. australis), northern fur seals (Callorhinus ursinus), and probably several other species. Sea otters(Enhydra lutris, Mustelidae) are commonly killed by white sharks off California, but have yet to be found as stomachcontents. Mammalian carrion from slaughterhouses and other sources, including mutton, pig, horse, dog, cattle, and rarelyhuman, has been found in the white shark’s stomach also. Terrestrial mammals probably do not figure as a significant part ofthe white shark’s diet.Larger white sharks above 3 m long tend to prey more readily on marine mammals than smaller sharks below 2 m long whichfeed more readily on bony fishes and sharks (although the young of smaller pinnipeds can be taken by small white sharks).This and other observations have lead to the belief that large white sharks are virtually dependent on pinnipeds for prey.However, large white sharks are not restricted to pinniped prey (even in areas with pinniped colonies and abundant seals),but also catch large teleost fishes, sharks and rays, birds, dolphins and marine reptiles, and are presumably capable ofsubsisting on such other small to large prey, in areas where seals are uncommon or absent (Mediterranean Sea, SpencerGulf in South Australia, Brazil). One 4.4 m specimen from the USA (Washington State) had 150 crabs (Cancridae) as well assalmon, hake, rockfish and seal hides in its stomach (LeMier, 1951). Pinnipeds may be especially important prey for whitesharks where they occur together, especially at seal colonies where pinnipeds are highly vulnerable. An observationalproblem is that seal predation by white sharks at concentrated seal colonies can be more easily studied than theirinteractions with other prey items. Rather than comprising a simple growth-related switch from fish to mammal prey as someauthors have assumed, the prey spectrum of the white shark is more likely to increase with growth in maximum prey size andin diversity. Only larger white sharks are apparently able or motivated to kill large pinnipeds, dolphins, bony fish, sharks, raysand turtles while retaining the ability to capture smaller prey.Recent observations and filming of white sharks at Dyer Island, South Africa, suggest that larger animals are anything butinept, clumsy predators. The slow cruise speed and deliberate investigation of boats and baits by large white sharks beliestheir high dash speed and ability to chase, overhaul, and kill fast, agile prey such as Cape fur seals, sometimes after multiplejumps. Large, live, active prey may be taken by a sudden, swift rush at high speed, sometimes from below but also at thesurface or even after jumping into the air. The white shark is inconspicuously coloured when viewed from above and below,and may take advantage of its cryptic coloration to stalk potential prey (as with apical terrestrial predators), but it also willcruise through Cape fur seal rafts and suddenly attack a selected victim. If the shark misses its prey, a fast chase (includingjumping) may ensue. Attachment of sonic tags (Strong et al, 1992) and self-contained remote cameras (‘crittercams’, I.Fergusson and G. Marshall, pers. comm., and supplied footage) to white sharks show slow patrolling just below the surfaceand near the bottom, punctuated by occasional fast dashes (apparently after prey), with little time spent in midwater.Invertebrate prey includes squid (Loliginidae), abalone (Haliotis) and other gastropods, bivalves, and crabs (Cancridae).Inedible garbage is occasionally taken from the stomachs of white sharks, but apparently this species is not fond ofswallowing oddities as does the tiger shark.Much speculation has occurred on the predatory behaviour of white sharks, particularly in the context of biting people. Thewhite shark is clearly capable of inflicting mortal wounds on a human with a single bite, but often merely clamps lightly onto adiver or swimmer without completing its bite and releases them after a short time. Such inhibited biting behaviour has beeninterpreted as a predation tactic (‘bite and spit’) of an inept, ambush predator to avoid injury from prey, but is unlikelybecause of the nature of full-fledged white shark predation bouts on large active, potentially injurious prey such as seals. Awhite shark generally delivers a powerful bite (or bites) to immobilise, incapacitate or kill the prey item outright, then maycontinue biting while feeding, may resume biting after a short to long interval, or may leave without consuming more of the

Sharks of the World, Vol. 2 105victim. The presence of other white sharks near a kill may influence the predator’s activities via social interactions. An‘exsanguination’ predation tactic has been suggested for white sharks (Klimley, 1994), in which the shark bites to kill a victimby bleeding it to death. However, white shark bites often cause severe or mortal injury (including decapitation and removal oflimbs) apart from blood loss. The predation tactic used by white sharks on large, active prey is more likely the rapid inflictionof ‘massive trauma’ to halt and kill its victim, which also begins the feeding process (somewhat as in pack-hunting terrestrialcarnivores that run down their ungulate prey and eat them to death). Smaller prey items including neonate Cape fur sealsmay be swallowed whole by large white sharks without powerful biting or chasing (sometimes after slowly following them atthe surface), and this may be likewise with large sharks eating small bony fish, crabs, and other small, defenceless prey.Non-feeding, inhibited bites on people may be agonistic, such as possibly happens in contact encounters among whitesharks, or alternatively exploration, displacement (displaced aggression), or even play. Inhibited bites have been explainedas ‘mistaken identity’ predation bouts, in which the shark bites its victim after mistaking it for a seal or turtle and then abortsits activity after biting. Observation on free-ranging white sharks suggests that white sharks are highly selective visualpredators and may be readily able to distinguish prey from divers or surfers and will react accordingly. Mistakes are possible,however, when visibility is impaired. Full-fledged predation bouts in which the white shark repeatedly bites and dismembersa human victim have been reported, but fortunately these are extremely rare compared to inhibited bites, and are very rarecompared to episodes of human predation by large terrestrial carnivores, particularly tigers and the largest macropredatorycrocodilians.Ironically, Homo sapiens is the chief known predator and cause of mortality to white sharks, through targeted and bycatchfisheries. A large orca (Orcinus orca) was recently seen to kill a white shark off the Farallon Islands, California, UnitedStates, but little is otherwise known of the interactions between these apical predators. Orcas and white sharks broadlyoverlap in distribution and take much the same prey (except for large baleen whales occasionally killed by orcas), butcomplement each other ecologically and possibly displace each other in different areas. Colder seas near the poles tend tohave more killer whales, cool to warm-temperate seas more white sharks, while both can occur in the tropics. Thebehavioural and ecological relationships of killer whales and white sharks are little understood. Large pinnipeds and otherspecies of large, macropredatory sharks are potential predators or sources of injuries to white sharks, but without muchevidence except occasional seal bite-marks on sharks. White sharks will also cut and bite one another in possibledominance-related aggressive interactions, and adult male white sharks may bite females during courtship. The injuries areoften minimal and apparently inhibited and it is not known what the contribution of such injuries are to white shark mortality.Nothing is known of white shark bacterial or viral diseases, or if common parasites such as the large copepods on themaxillary valve of white sharks or the intestinal cestode fauna are a source of problems for them. Sometimes large whitesharks are found dead on beaches with no obvious external or internal injuries.Size: Maximum total length to about 6 m, and possibly to 640 cm or more; the largest free-swimming individuals commonlycaptured are between 500 and 580 cm (mostly adult females). Records of white sharks 6.4 to 7 or even 9 m long are difficultor impossible to verify. A much quoted record of an 11 m (36 ft) shark from Australia is erroneous and based on a jaw from ashark 4.9 to 5.5 m long in the British Museum of Natural History (Natural History Museum). Size at birth is assumed as beingbetween 109 and about 165 cm, with term foetuses at 100 to 165 cm and free-living young down to 109 to 129 cm. Malewhite sharks mature at a smaller size than females and reach a smaller maximum size. Size at maturity for males is about350 to 410 cm, with adolescence roughly between 250 and 400+ cm, and with small but fully adult males at 356, 368 and380 cm; maximum length for males is between 501 and at least 517 cm and possibly 550 cm. Females mature somewherebetween 400 and 500 cm, with females 441, 470 and 490 cm being immature or early adolescent (Florida, United States,and South Africa) and adults reported at 420 to 580 cm; maximum size of females possibly about 6 m.Several length-weight equations are available for the white shark. See Mollet and Cailliet (1996) for a detailed review oflength-weight methodology.Compagno (1984): W(kg) = 4.34 x 10 -6 TL(cm) 3.14(n = 98, TL = 127 to 554 cm, mostly from California)Tricas and McCosker (1984): W(kg) = 3.8 x 10 -6 x TL(cm) 3.15 (n = 127)Casey and Pratt (1985): W(kg) = 4.804 x 10 -6 x TL (cm) 3.095(n = 200, from the western North Atlantic)Cliff, Dudley and Davis (1989): W(kg) = 1.84 x 10 -5 x PCL(cm) 2.97Cliff, Dudley and Jury (1996): W(kg) = 2.14 x 10 -5 x PCL(cm) 2.944(n = 309, from South Africa)(n = 383, from South Africa)Kohler, Casey and Turner (1995): W(kg) = 7.5763 x 10 -6 x FL(cm) 3.0848 (n = 125)where FL(cm) = 0.9442 x TL - 5.7441 (n = 112, from the western North Atlantic)Compagno (update from 1984): W(kg) = 3.026 x 10 -6 x TL(cm) 3.188(n = 156, from California and South Africa).Interest to Fisheries and Human Impact: The importance of the white shark as a fisheries species is limited because of itslow abundance wherever it occurs, which restricts targeted commercial fisheries for conventional bulk fisheries productssuch as meat or liver oil. However, the high value of its jaws, teeth and fins makes it a viable target of small-scale targetedcommercial fisheries as well as an added value to bycatch. It is mostly caught as a bycatch of fisheries for other sharks andother marine organisms (including bony fishes and cephalopods), by many forms of gear utilized by modern marine fisheries

106 FAO Species Catalogue for Fishery Purposes No. 1including longlines, specialized heavy line gear, rod-and-reel, fixed bottom gill nets, floating inshore gill nets, pelagic gillnets, fish traps, herring weirs, trammel nets, harpoons, bottom and pelagic trawls, and purse seines. Its tendency toinvestigate human activities (including fishing operations) and to scavenge from fishing gear, as well as the value of its jawsand fins, makes it very vulnerable to being killed and captured despite its size and strength. It is subject to targeted sportsfisheries for game-fishing records and trophy jaws because of its great size, notoriety, and powerful resistance to capture. Itis also targeted by small-scale and erratic commercial fisheries in several countries.The meat of the white shark is or has been utilized fresh, fresh-frozen, dried-salted, and smoked for human consumption,although the extremely high mercury content of the meat limits its utility. The flesh has been used in traditional medicine inSouth Africa. White shark meat has been sold as ‘shark’ in California, but in the 1980s at least one market got higher valuesby marketing it as white shark meat. The liver of the white shark has been extracted for vitamin oil. Presumably white sharkcartilage is processed for medicinals but the writer has no evidence of it. White shark carcasses have been processed forfishmeal, but also frozen or preserved whole for exhibit in oceanaria and museums. White shark specimens are also castand modelled whole for oceanarium and museum exhibits and for trophies. The skin of the white shark has been utilized forleather, although rarely at present.White shark teeth and jaws are used for trophies, decorations and collectibles. Properly prepared white shark jaws maybring an inflated price, essentially what the upscale international private collector’s market will bear, with greatest value forthe jaws and teeth of large sharks over 5 m long. In South Africa, offers of US$20 000 to 50 000 have been made for whiteshark jaws, and US$600 to 800 for individual teeth.Apart from their size, white shark fins are boosted in value because of notoriety. A fin set from a large white shark may bevalued at over US$1 000. Unfortunately, as with rhino horns and elephant tusks the high value of white shark productsencourages poaching, clandestine trade, and flouting of protective laws.Live white sharks have been avidly sought by public aquaria and oceanaria for exhibits, but capture trauma usually insuresthat the sharks perish after a day or a few days in captivity. White sharks were mostly caught for display during the 1970s and1980s, but seldom are taken at present.The white shark has for the last few decades been the subject of commercial underwater cage-diving operations in SouthAustralia allowing sport SCUBA and snorkel divers to view and film white sharks, as well as intensive and sometimesoral-obsessive filming of these sharks by professional film crews. This activity also started in South Africa in the early 1990s,continues unabated at present, and has engendered some problems for the sharks and for researchers trying to study them.Regulation of capture and ecotouristic access to white sharks was seriously debated in Australia and South Africa, andlicensing and guidelines have been imposed to limit access to white shark sites and methods of attracting white sharks toboats. A flurry of shark-bite incidents off South Africa in the late 1990s spawned a high-profile media-hyped controversy onthe assumption that exposure of white sharks to cage diving in the Western Cape somehow caused them to bite peopleelsewhere.The white shark is feared by many people as the most ‘dangerous’ living shark; more instances of white sharks bitingswimmers, divers, surfers, and boats have been reported than for any other shark. Although much has been made of whiteshark encounters with people and boats in the popular news and entertainment media, including the JAWS motion picturesthat established the white shark as a Hollywood science-fiction monster, the white shark is not very ‘dangerous’ whencompared to other causes of injuries and fatalities to people. Most white shark bites are nonfatal and may not be predatory(see above). The rate of white shark bite incidents averaged about three per year worldwide between 1952 and 1992 (range0 to 8, with an 18% fatality rate) and a slowly increasing trend from about 1.5 per year between 1952 and 1962 to 5.3 per yearbetween 1983 and 1992 (data in part from Ellis and McCosker, 1992; see also Burgess and Callahan, 1996). During the lastdecade (1990 through 1999) unprovoked incidents averaged 6.9 per year (69, range 0 to 12 per year, with a 16% fatalityrate), with most occurring off the United States and South Africa (G. Burgess, International Shark Attack File, pers. comm.).This is far less than drownings, diving accidents, automobile accidents, deaths from lightning strokes, injuries or deaths fromterrestrial animals including tigers, large crocodilians, and even domestic livestock, or other calamities that afflicted humansin the countries where white shark incidents occurred.Most injuries and fatalities from white shark encounters have occurred off California (United States), southern Australia,New Zealand, South Africa, and Japan, but about 80% of reported shark biting incidents have occurred in the tropics, wherewhite sharks are rare or uncommon but where large carcharhinid sharks predominate. There might very well be tropical andwarm-temperate carcharhinoids, particularly the tiger and bull sharks, that may be responsible for more shark bite incidentsthan the white shark, but this remains to be verified.Several surfers and paddleboarders have been bitten by white sharks or knocked off their boards. The sharks havesometimes destroyed the boards or held, shook, and released the boards without harming the surfers. Boats may be bitten,particularly if boaters provoke the sharks by landing fish out of their grasp or try to ram the sharks. For some reason FalseBay, South Africa, was the site of more boat incidents involving white sharks than all other places combined between 1940and 1980, although hardly since then. White sharks have on a few occasions persistently bitten at boats until they sank, andin a few instances leapt into a boat or punched through the hull like a torpedo.Despite the frightful, JAWS-inflated reputation of the white shark there is accumulating evidence that the presence of a whiteshark near people (especially divers) does not necessarily mean that the shark will automatically bite (much less kill and eat)them. Numerous incidents have been reported where these sharks swim up to divers as close as a few feet and depart

Sharks of the World, Vol. 2 107without biting or without displaying possible agonistic behaviours such as gape or hunch, and these have been followed upby deliberate encounters by film-makers with white sharks outside shark cages without adverse affects. Numerous close,deliberate, and systematic encounters involving divers without a cage in proximity to numerous white sharks ranging in sizefrom about 2 to over 5 m long have shown very little agonistic behaviour by the sharks, although they were inquisitive, weresometimes present in groups, and often came very close to the diver (M. Marks pers. comm. and videos). This is definitelynot recommended for ordinary divers seeking recreational thrills, and should be seen as deliberate and calculated risk insupport of science comparable to behavioural observation of large, potentially harmful terrestrial carnivores and largeungulates.World catches of white sharks from all causes are difficult to estimate. Compagno (1984) noted that off California 10 to 20 ormore white sharks were killed each year as a bycatch of various fisheries, vs. 0.13 humans per year killed by white sharks.These figures were over a period of about three decades up to the early 1980s, and protection of the white shark in the1990s may have reduced the catch since then.Conservationists have been concerned with the vulnerability of the white shark to possible extinction, and it is currentlyconsidered a vulnerable species in the IUCN Red List. This seems warranted given that this species has a relatively lowintrinsic rebound potential (a relative measure of the ability to recover from fishing; Smith, Au and Show, 1998). Threats tothe white shark include: 1. Targeted commercial and sports fisheries for jaws, fins and game fish records. 2. Anti-sharkmeasures to protect beaches. 3. Ad-hoc media-fanned campaigns to kill white sharks after a biting incident occurs.4. Long-term increases in use and efficiency of commercial and artisanal fishing gear that have an impact on white sharks.5. Degradation of inshore habitats used by white sharks for pupping and nurseries. 6. Increased demand for shark productsin general, including vastly inflated values of white shark jaws, fins and teeth. 7. The tendency of white sharks to investigatehuman activities, which can be fatal to them. 8. Lack of knowledge on many crucial aspects of white shark biology, andlimited interest on investigating them. 9. The negative JAWS image, which invites neglect, killing of sharks, and disregard ofconservation measures. 10. Slow maturation and low fecundity. 11. Proximity of concentrations of white sharks tohighly-developed coastal areas such as those of California, the Mid-Atlantic Bight, and the Mediterranean Sea. 12.Inadequate catch data in most places where the white shark occurs.Numbers of white sharks have been thought to be declining off Australia due to sports and commercial fisheries andanti-shark nets. The white shark is currently protected in Australia (nationally and statewise), South Africa, Namibia, Israel,Malta and the United States (statewise off California and Florida and with directed fisheries prohibited off all coasts).Protective laws are strict, but loopholes and inadequate enforcement may cause problems including promoting a blackmarket for high-value white shark products. The United States and Australia proposed the white shark for CITES I and IIlisting at the 2000 CITES Conference to ban trading in white shark parts, but the proposals were not approved. Thesecountries are currently proposing the white shark for a CITES III listing while Australia has developed a comprehensive andmultidisciplinary recovery plan for white sharks in its waters (Environment Australia, 2000b).Local Names: Sarda (Canary Islands); Jaquetón de ley (Cuba); Uptail, Tommy, White death shark, White death, Greatwhite death, Death shark, White pointer (Australia); Great white shark, White shark (AFS), Maneater or man-eater shark,(England and USA); Ami, Lamea, Lamie, Lameo, Le Carcharodonte lamie, Le grand requin, Pei can, Requin blanc (France);Menschen fresser, Menchenhai, Merviel fras, Weisshai (Germany); Ca mari, Marraco, Salroig, Salproig, Salproix, Taburo,Tiburo, Tiburón blanco, Tauró blanc (Spain); Tubarâo branco, White shark (Azores); Carcarodonte, Cagnia, Cagnescagrande, Cagnia, Caniscu, Carcarodonte lamia, Carcarodonte di rondelet, Damiano [sic] or Demon shark, Imbestinu, Lamia,Mastinu feru, Mangia alice or Anchovy-eater, Pesce cane, Pesca can, Pescecane, Pesce can grande, Pesciu can, Picibistinu, Pisci cani or Can grossu, Pisci mastinu, Squalo bianco, Tunnu palamitu di funnu or Tuna shark (Italy); Hohojirozame,Hitokiuzame, Oshirozame (Japan); Psina ljudozdera (Yugoslavia); Gab doll, Kelb Il-bahar, Kelb-il-bahar abjad, Hutatax-xmara (Malta); Taniwha, Mango-tuatini, Hare hongi (New Zealand); Haa skieding (Norway); Tabarao (Portugal); Gench,Kersch (Red Sea); Blue pointer, White shark, Great white shark, Springhaai or Jumping shark, Witdoodshaai, Withaai,Cowshark, Mudshark, Sarda (South Africa); Niuhi (Hawaiian Islands); Tanifa (Samoa).Literature: Linnaeus (1758); Müller and Henle (1839); Smith (1849); Garman (1913); Barnard (1925, 1937); Fowler (1936,1941); Springer (1939); Whitley (1940); Bonham (1942); Bigelow and Schroeder (1948); Smith (1949, 1951); LeMier (1951);Herre (1953); Schultz et al. (1954); Strasburg (1958); Farquhar (1963); Smith and Smith (1963); Garrick and Schultz (1963);Limbaugh (1963); Stead (1963); Follett (1966); Squire (1967); Case (1968); Carey and Teal (1969); Mundus and Wisner(1971); Arnold (1972); Randall (1973, 1987); Springer (1973); Bass, D’Aubrey and Kistnasamy (1975a); Ellis (1976); Allenand Allen (1978); Wallett (1978); Penrith (1978); Ames and Morejohn (1980); Ainley et al. (1981, 1985); Miller and Collier(1981); Carey (1982, 1990); Carey et al. (1982, 1985); le Boeuf, Riedman, and Keyes (1982); Pratt, Casey and Conklin(1982); Brodie and Beck (1983); Compagno (1984, 1990a, b, d, 1991); Engaña and McCosker (1984); Quero (1984); Tricasand McCosker (1984); Block and Carey (1985); Cailliet et al. (1985); Pratt and Casey (1985); Klimley (1985); Taylor (1985);Tricas (1985); Tricas and McCosker (1985); McCosker (1985); Bass (1986); deSilva (1986); Corkeron, Morris and Bryden(1987); Nakano and Nakaya (1987); Stafford-Deitsch (1987); Randall, Randall and Compagno (1988); Uchida et al. (1988);Paxton et al. (1989); Cliff, Dudley and Davis (1989); Cockcroft, Cliff and Ross (1989); Paulin et al. (1989); Fulgosi (1990);Strong (1990, 1996); Springer (1990); Cousteau and Richards (1991); Ellis and McCosker (1991); Long (1991, 1996); Bruce(1992, 1995); Klimley et al. (1992); Strong et al. (1992, 1996); Francis and Randall (1993): Hanan, Holts and Coan (1993);Michael (1993); Compagno and Fergusson (1994); Klimley (1994); Seret (1994); Last and Stevens (1994); Bonfil (1994);Fergusson (1994, 1996); Nakaya (1994, 1996); Anderson and Goldman (1996); Burgess and Callahan (1996); Cliff, Dudleyand Jury (1996); Collier, Marks and Warner (1996); Goldman et al. (1996); Francis (1996); Gadig and Rosa (1996); Klimley,Pyle and Anderson (1996); Long and Jones (1996); Long and Waggoner (1996); Long et al. (1996); Mollet and Caillietclick for next page

click for previous page108 FAO Species Catalogue for Fishery Purposes No. 1(1996); Mollet et al. (1996); Pratt (1996); Strong (1996); Uchida et al. (1996); Goldman (1997); Smale and Heemstra (1997);Compagno, Marks and Fergusson (1997); Santos, Porteiro and Barreiros (1997); McEachran and Fechhelm (1998); Smith,Au and Show (1998); Cliff et al. (2000); Environment Australia (2000b); Fergusson, Marks and Compagno (2000); D. Ainley(pers. comm.); R. Alexander (pers. comm.); D. Allen (pers. comm.); G. Burgess (pers. comm.); G. Cailliet (pers. comm.); S.Cook (pers. comm.); D. Ebert (pers. comm.); A.P. Klimley (pers. comm.); R. Lea (pers. comm.); M. Marks (pers. comm.); S.Smith (pers. comm.); I. Fergusson (pers. comm.); G. Zorzi (pers. comm.).Isurus Rafinesque, 1810Genus: Isurus Rafinesque, 1810a, Caratt. gen. sp. anim. piant. Sicilia, Palermo, pt. 1: 11.Type Species: Isurus oxyrinchus Rafinesque, 1810, by monotypy.Number of Recognized Species: 2.Synonyms: Genus Oxyrhina Agassiz, 1838: 86, pls. 33-34. Type species: “Lamna oxyrhina Cuvier and Valenciennes,MS.” by absolute tautonymy, a junior synonym of Isurus oxyrinchus Rafinesque, 1810. Genus Oxyrrhina Bonaparte,1846: 17. Type species: Oxyrhina gomphodon Müller and Henle, 1839, a junior synonym of Isurus oxyrinchusRafinesque, 1810. Genus Plectrostoma Gistel, 1848: 10. Replacement name for Oxyrhina Agassiz, 1835, and therebytaking the same type species, Lamna oxyrhina Cuvier and Valenciennes, 1838, a junior synonym of Isurus oxyrinchusRafinesque, 1810. Genus Isuropsis Gill, 1862b: 397. Type species: Oxyrhina glauca Müller and Henle, 1839, by originaldesignation. Genus Plectrosoma Bigelow and Schroeder, 1948: 123 (error for Plectrostoma Gistel, 1848). GenusLamiostoma Glikman, 1964: 105. Type species: Lamiostoma belyaevi Glikman, 1964, by original designation.Diagnostic Features: Snout bluntly to acutely conical. Eyes large, 1.3 to 3.1% of total length. Nostrils situated adjacent tohead rim in ventral view. Mouth width 0.9 to 1.5 times its length. Anterior teeth enlarged; anterior and intermediate teeth notcompressed but upper lateral teeth more or less compressed, teeth not forming a continuous cutting edge; intermediateteeth very small and less than half height of adjacent anteriors, with normal cusps directed posterodistally or nearly vertical;second lower anterior teeth greatly enlarged and taller than second upper anterior tooth; total tooth count 43 to 54; roots ofanterior teeth deeply arched, with root lobes narrow and elongated; lateral cusplets absent from teeth at all stages; teethsmooth-edged; cusps of anterior teeth weakly to strongly flexed. Body usually fairly slender. First dorsal-fin origin usuallybehind the pectoral-fin free rear tips. Anal-fin origin under second dorsal-fin midbase or slightly posterior to second dorsal-fininsertion. Secondary caudal keels absent or present and weak. Total vertebral count 182 to 197. Cranium with rostralcartilages neither swollen nor hypercalcified. Intestinal valve count 47 to 54. Length of adults 2.0 to 4.2 m. Usually a blackaxillary spot at pectoral-fin insertions; pectoral-fin tips usually abruptly black on their ventral surfaces.Local Names: Makos, Mako sharks, Mackerel sharks, Awozame-zoku (Japan).Remarks: The systematics of makos (genus Isurus) was formerly chaotic, with a few regional species recognized ongrowth changes within a single species, Isurus oxyrinchus (Garman, 1913; Fowler, 1941; Bigelow and Schroeder, 1948;Smith, 1957). Guitart (1966) described a second, highly distinctive macroceanic mako, I. paucus from Cuba, which Garrick(1967) independently described as I. alatus from the central Pacific. Garrick (1967) sorted out the growth change problemsand his arrangement and synonymy for I. oxyrinchus were followed by Bass, D’Aubrey and Kistnasamy (1975a), Pinchuk(1983), Compagno (1984), and Moreno and Morón (1992b), who recognized two living species, I. oxyrinchus andI. paucus. Moreno and Morón presented detailed comparisons of eastern Atlantic makos from extensive fisheries samplesand confirmed the differentiation of I. oxyrinchus and I. paucus on morphometry, external morphology including coloration,and dentitional characters. They noted, however, that I. oxyrinchus specimens caught around the Azores closely agreedwith other eastern Atlantic specimens in morphometrics, morphology and dentition, but had a dusky snout and mottled orall-dark underside as in I. paucus. They suggested that the Azores mako represented a distinct, localized population or‘population variant’ of I. oxyrinchus. This is plausible but needs further research, including molecular studies, and is ofsome urgency as the Azores mako is possibly declining due to overfishing. The Azores mako may have an analogy withinthe genus Lamna, in which the North Pacific L. ditropis differs from the North Atlantic L. nasus in having a dusky or mottledunderside, but Southern Hemisphere ‘black-faced’ L. nasus have similar dark undersides and have been mistaken forL. ditropis. Compagno (1990b) noted chondrocranial differences between Isurus oxyrinchus and I. paucus.

Sharks of the World, Vol. 2 109Key to Species:1a. Snout usually acutely pointed; cusps of upper andlower anterior teeth flexed, with tips reversed;pectoral fins considerably shorter than head,relatively narrow-tipped in young and acutelypointed in adults (Fig. 72); underside of snout andmouth white in adults (dusky in Azores’ ‘marrajocriollo’). . . . . . . . . . . . . . . . . . Isurus oxyrinchus1b. Snout narrowly to bluntly pointed, usually notacute; cusps of upper and lower anterior teethstraighter, with tips not reversed; pectoral finsabout as long as head or longer (Fig. 73),relatively broad-tipped in young and adults;underside of snout and mouth dusky in adults. . . . . . . . . . . . . . . . . . . . . . . . Isurus paucusFig. 72 Isurus oxyrinchusFig. 73 Isurus paucusIsurus oxyrinchus Rafinesque, 1810 Fig. 74Isurus oxyrinchus Rafinesque, 1810a, Caratt. gen. sp. anim. piant. Sicilia, Palermo, pt. 1: 12, pl. 13, fig. 1. Also Rafinesque,1810b, Indice Ittiol. Sicil.: 45. Holotype unknown, type locality Sicily, Mediterranean Sea. Variant spellings include Lamiaoxyrhincus Bory de St. Vincent, 1829, and Isurus oxyrhynchus Jordan and Evermann, 1896.Synonyms: Isurus spallanzanii Rafinesque, 1810b: 45, 60. Type locality: Sicily. No types known according to Eschmeyer(1998: CD-ROM). Squalus (Lamna) cepedii Lesson, 1830: 93. Holotype: 1.83 m individual, Equatorial Atlantic, 6°S, 27°W;according to Eschmeyer (1998: CD-ROM) the whereabouts of this specimen is unknown. Lamna oxyrhina Cuvier andValenciennes, in Agassiz, 1838: 86. Manuscript name only cited by Agassiz, but without a species description. Typesuncertain: Agassiz mentioned a set of jaws in the collection of the Museum National d’Histoire Naturelle, Paris, withoutspecific data. A tooth set from a mako was illustrated by him (Agassiz, 1835, pl. G, fig. 2, as Lamna) and may be from theParis specimen. Oxyrhina gomphodon Müller and Henle, 1839: 68, pl. 28. Holotype, Zoologisches Museum, Museum fürNaturkunde der Humboldt-Universität, Berlin, ZMB 4528, a stuffed adult male 1 650 mm +, without head, jaws separateaccording to Paepke and Schmidt (1988: 163), from “Ocean”. Oxyrhina glauca Müller and Henle, 1839: 69, pl. 29.Syntypes, three stuffed specimens, Rijksmuseum van Natuurlijke Historie, Leiden, RMNH 2529, 1 020 mm TL male (alectotype, designated by Garrick, 1967: 675), RMNH 2533, 750 mm TL male, RMNH 2587, 660 mm TL male, and two jaws ofuncertain deposition, possibly Berlin (not mentioned in Paepke and Schmidt, 1988). Type locality originally stated as Java,but actually Japan according to Garrick (1967). Lamna latro Owen, 1853: 96. Locality unknown. Types? Isuropsis dekayiGill, 1862: 409. Based on Lamna punctata DeKay, 1839, not Oxyrhina daekayi Gill, 1862b, (= Lamna nasus). Carchariastigris Atwood, 1865: 81. No distinguishing features; also Atwood, 1869: 268. Types: None, according to Eschmeyer (1998:CD-ROM), from Gulf of Mexico and Massachusetts, Provincetown, Cape Cod. Lamna guentheri Murray, 1884: 349. Typelocality, Kurrachee (= Karachi), Pakistan. Original spelling Lamna güntheri. Holotype: a 2 580 mm specimen according toGarrick (1967: 665-667). Disposition of holotype uncertain, presumably in the British Museum (Natural History); Garrick (loc.cit.) and Eschmeyer (1998: CD-ROM) had no information. Lamna huidobrii Philippi, 1887: 548, pl. 3, fig. 1. Holotype:2.90 m female (possibly stuffed) in collection of Universidad de Chile, Departamento de Biología Celular y Genética,Santiago, Chile (Eschmeyer, 1998: CD-ROM), possibly lost (S. Kato, pers. comm.), type locality, Santiago Province, Chile.Isurus mako Whitley, 1929: 101. A new name based on the account of I. glaucus by Phillipps, 1932: 268, off Manaukau Bar,Auckland, New Zealand. No types according to Eschmeyer (1998: CD-ROM). Isurus bideni Phillipps, 1932: 227, fig. 2.According to Phillipps (loc. cit.) and Eschmeyer (1998: CD-ROM), the holotype is a South African specimen in the AustralianMuseum, Sydney, AMS IA.4311 (jaws and photographs) sent to Mr G.P. Whitley by Mr C. Leo Biden of Cape Town andpossibly from the Western Cape Province, South Africa. Isurus tigris africanus Smith, 1957: 96, pl. 1, fig. 1. Holotype:J.L.B. Smith Institute of Ichthyology, RUSI 426, jaws from 2 540 mm TL adult male. Type locality, Algoa Bay, South Africa.Other Combinations: Isurus or Isuropsis glaucus (Müller and Henle, 1839), Isurus guentheri (Murray, 1884).

110 FAO Species Catalogue for Fishery Purposes No. 1FAO Names: En - Shortfin mako; Fr - Taupe bleu; Sp - Marrajo dientuso.UPPER AND LOWER TEETH OF LEFT SIDEUNDERSIDE OF HEADFig. 74 Isurus oxyrinchusField Marks: Spindle-shaped body, long, acutely conical snout, large blade-like teeth without cusplets or serrations,pectoral fins rather narrow-tipped and with anterior margins less than head length, large first dorsal fin and minute, pivotingsecond dorsal and anal fins, strong keels on caudal peduncle, no secondary keels on caudal base, crescentic caudal fin,ventral surface of body usually white.Diagnostic Features: Snout acutely pointed. Eyes relatively small. Mouth U-shaped. Lower anterior teeth stronglyprotruding and horizontal on jaws even when mouth is closed; anterior teeth with narrow, more oblique cusps with flexedtips; cusps of first upper anterior teeth with incomplete cutting edges; intermediate teeth with curved oblique distally hookedcusps. Body moderately slender but more fusiform than in Isurus paucus. Pectoral fins semifalcate and with tips relativelynarrow, anterior margins about 16 to 22% of total length and shorter than head length. Origin of first dorsal fin over or justbehind the pectoral free rear tip; first dorsal-fin apex broadly rounded in young but more angular and narrowly rounded inlarge juveniles and adults; first dorsal-fin height greater than base length in large individuals but equal or smaller in youngbelow 185 cm. Total vertebral count 182 to 195, mostly below 190. Colour (except Azores ‘marrajo criollo’): dorsolateralcoloration brilliant blue or purplish in life, white below underside of snout in young and adults; dark colour of head partiallycovering gill septa, lower part of second and third gill septa white; dark colour of flanks not extending ventrally ontoabdomen; pelvic fins dark on anterior halves, white on posterior halves, undersides white and sometimes with thin darkmargin; first dorsal fin with pale centre (more obvious in young than adults); anal fin dark only on anterior half and white onposterior half.Distribution: Coastal and oceanic, circumglobal in all temperate and tropical seas. Western Atlantic: Gulf of Maine tosouthern Brazil and possibly northern Argentina, including Bermuda, Gulf of Mexico and Caribbean. Eastern Atlantic:Norway, British Isles and Mediterranean to Morocco, Azores, Western Sahara, Mauritania, Senegal, Côte d’Ivoire, Ghana,southern Angola, probably Namibia, and South Africa (west coast). Indo-West Pacific: South Africa (east coast),Mozambique, Madagascar, Mauritius and Kenya north to Red Sea and east to Maldives, Iran, Oman, Pakistan, India,Indonesia, Viet Nam, China, Taiwan (Province of China), North Korea, South Korea, Japan, Russia (Primorskiyi Kray),Australia (all states and entire coast except for Arafura Sea, Gulf of Carpentaria and Torres Strait), New Zealand (includingNorfolk Island), New Caledonia, Fiji. Central Pacific: From south of Aleutian Islands to Society Islands, including HawaiianIslands. Eastern Pacific: USA (Washington and California) south to Mexico, Costa Rica, Ecuador, Peru and central Chile.

Sharks of the World, Vol. 2 111Habitat: The shortfin mako is a common, extremely active, offshore littoral and epipelagic species found in tropical andwarm-temperate seas but seldom occurring in waters below 16°C. This shark occurs from the surface down to at least500 m. It occurs well offshore but penetrates the inshore littoral just off the surf zone in places such as parts ofKwaZulu-Natal, South Africa, where the continental shelves are very narrow. Shark meshing data off South Africa suggeststhat this species prefers clear water to turbid water and is caught at a range of water temperatures from 17 to 22°C. In thewestern North Atlantic it prefers a similar range of temperatures, and only moves onto the continental shelf when surfacetemperatures exceed 17°C. In the eastern North Pacific juveniles range into southern California waters and tend to stay nearthe surface, with little tendency to descend into cold subsurface waters, and use these offshore continental waters asnursery areas. Telemetered small sharks in this area tended to stay near the surface above 20 m depth and in waters 20 to21°C, and mostly avoided the thermocline and cold deeper waters (Holts and Bedford, 1992).Biology: The peregrine falcon of the shark world, the shortfin mako may be the fastest shark and one of the swiftest andmost active fishes. It is famed as a jumper, leaping several times its length from the water, and is capable of extreme burstsof speed when hooked and in pursuit of prey. Sonic telemetry of small (1.7 to 1.8 m and 32 to 36 kg) makos off southernCalifornia showed a relatively slow cruise speed (Holts and Bedford, 1992) between 1.3 and 2.6 km/h, with short increasesup to 4.5 and 5.6 km/h and slowing to less than 1 km/h at night. No fast dashes were recorded during the periods monitored.Similar speeds were recorded over a track of 280 km off Florida by a larger (about 180 kg) shortfin mako (Casey and Kohler,1992).Shortfin makos are endothermic and maintain higher temperatures than surrounding water temperatures in their bodymusculature, brains, eyes and viscera with countercurrent vascular heat exchangers (Carey and Teal, 1969; Carey et al.,1972; Carey, 1982, 1992; Block and Carey, 1985). Body muscle may run 1 to 10°C higher than ambient temperature. Centraltemperatures in the body musculature of the shortfin mako are highest relative to water temperature in cooler water(between 19 and 25° in water 15°C), but approach ambient in warm water (27 to 29° in water 27°C). Endothermy isparticularly advantageous in maintaining elevated swimming, visual, central nervous system and digestive functions whenthe makos are in cooler waters in higher latitudes or when diving below the thermocline in warm waters.The shortfin mako is highly migratory and in the extreme northern and southern parts of its range, has a tendency to followmovements of warm water masses polewards in the summer. Off South Africa catches in the KwaZulu-Natal shark netsindicate inshore movements from deeper water over the continental slopes, which in the area is relatively close to shorebecause of the narrow shelves.Long-range movements of this shark are not well known and are being slowly elucidated by conventional tagging in theNorth Atlantic. Casey and Kohler (1992) reported on 2 459 shortfin makos tagged off the western North Atlantic coast of theUSA between 1962 and 1989, with a recapture rate of 9.4% to date. About 64% were recaptured within 500 km of where theywere tagged, but 13% travelled over 1 600 km with several recaptures in the Azores and one even travelling about 4 000 kmto Spain. Casey and Kohler suggested that makos may travel up the Gulf Stream and around the Sargasso Sea to return tothe East Coast of North America, with few crossing the Mid-Atlantic Ridge and reaching European waters. The authorshypothesized that the western North Atlantic makos form a separate population or stock to those in the eastern Atlanticalthough intermixing is possible as shown by crossings to the Azores and Europe. There are problems in understanding thepopulation structure of western North Atlantic makos because large adults, particularly females, are little-known in the areaalthough juveniles are abundant.

112 FAO Species Catalogue for Fishery Purposes No. 1This species is ovoviviparous and a uterine cannibal (oophagous), with 4 to 25 and possibly 30 young (mostly 10 to 18) in alitter, with larger females having larger litters. Sex ratios from beach-meshed sharks off KwaZulu-Natal, South Africa, variedmonthly, with male-female ratios ranging from 0.6:1 to 2.5:1 and with males more abundant than females for most of the yearexcept January and August; with a sample of 171 sharks, males were significantly more abundant than females with anoverall ratio of 1.4:1. Birth occurs mostly in late winter to midsummer in both hemispheres. Off KwaZulu-Natal birth occurredoffshore in late spring (November), and mating occurs in autumn (March to June). Estimates of gestation period vary, butrecent estimates suggest it may be 15 to 18 months within a reproductive cycle of three years (including a year breakbetween pregnancies; Mollet et al., 2000). Adult males show seasonal changes in gonadosomatic index (gonad weight as %of body weight), with males having the heaviest gonads in the summer and the lightest in winter. Gonadosomatic index ispositively correlated with hepatosomatic index (liver weight as % of body weight) in adult females, with females with largeovaries having larger livers than those with small ovaries.Makos may mature at a minimum age of 7 to 8 years old, based on yearly addition of growth rings on vertebral centra, withthe oldest known with 18 rings corresponding to at least 18 years at 321 cm, and an estimated maximum age of 45 years(Cailliet et al., 1983). Pratt and Casey (1983) assumed that two growth rings were formed each year on the vertebral centralof shortfin makos, though Cailliet et al. (1983) suggested that one ring per year was formed. Stillwell (1990) suggested thatmakos were faster-growing, and achieved adult size in 4.5 years for males and 7 years for females.The shortfin mako primarily feeds on other fishes, with a wide variety of prey recorded. Prey items are typically much smallerthan the mako, and off South Africa range between 10 and 35% of the length of the predator. However, Stillwell (1990)suggested that large makos shifted to large prey near their own size, with swordfish (Xiphias) weighing 180 kg or morebeing commonly taken by large and presumably adult makos (males about 136 kg and females 337 kg) in the western NorthAtlantic.The mako eats both pelagic and demersal bony fishes, including freshwater eels (Anguillidae), eel-pouts (Zoarcidae),anchovies (Engraulidae), menhaden, sardines and other herring (Clupeidae), grunts (Haemulidae), lancetfish(Alepisauridae), needlefish (Belonidae), sauries (Scomberesocidae), cod, ling, whiting and other cod-like fishes (Gadidae),hake (Merlucciidae), alfonsinos (Berycidae), croakers (Sciaenidae), mullet (Mugilidae), Australian salmon (Arripis),butterfishes (Stromateidae), bluefish (Pomatomus saltatrix, Pomatomidae), yellowtails, jack mackerel and other jacks(Carangidae), sea bass (Serranidae), porgies or sea breams (Sparidae), scabbardfish (Trichiuridae), snake mackerel(Gempylidae), swordfish (Xiphias gladius, Xiphiidae), sailfish (Istiophorus platypterus, Istiophoridae), mackerel, tuna,bonito, skipjack, and other scombrids (Scombridae), medusafishes (Centrolophidae), rockfish (Sebastes, Scorpaenidae),sea robins (Triglidae), and even seahorses (Syngnathidae), puffers (Tetraodontidae), porcupine fish (Diodontidae), andboxfish (Ostraciidae). In the western North Atlantic, bluefish are the most important food and comprise about 78% of theshortfin mako’s diet (Stillwell and Kohler, 1982), but the same species (called taylor in Australia) was not recorded frommakos off New South Wales (Stevens, 1984).Elasmobranchs taken by the shortfin mako include blue sharks (Prionace glauca), grey sharks (Carcharhinus obscurus,C. brevipinna, C. sealei and C. limbatus), milk sharks (Rhizoprionodon acutus), hammerheads (Sphyrna lewini and S.zygaena, skates (Rajidae), spotted eagle rays (Aetobatus narinari, Myliobatidae), and cownose rays (Rhinopteridae). OffKwaZulu-Natal, South Africa, small sharks less than 1.3 m long including the young of large species such as Carcharhinusobscurus are more important food items for the mako than large sharks and rays, which is a function possibly of highavailability as well as preference for smaller prey items. Off South Africa elasmobranchs were the most important prey,followed by teleosts and cephalopods (squid), but in the western North Atlantic and New South Wales teleosts were the mostimportant prey while elasmobranchs were virtually unrepresented.Cephalopods are important prey and include a variety of inshore and oceanic and deep-benthic squids including Loligo(Loliginidae), Ancistrocheirus (Ancistrocheiridae), Lycoteuthis (Lycoteuthidae), Octopoteuthis (Octopoteuthidae),Histioteuthis (Histioteuthidae), Gonatidae, and Illex and other ommastrephids (Ommastrephidae), as well as cuttlefish(Sepia, Sepiidae). Other food includes sea turtle heads (uncertain if these were scavenged or if the turtles were decapitatedby the makos), unidentified small cetacean material (including a pelagic dolphin), salps, isopods, penaeid shrimp, sponges,sargassum weed, and occasional stones and other detritus. Marine mammals are rarely reported in the diet of the shortfinmako and may be scavenged, but very large individuals occasionally catch small pelagic cetaceans (dolphins). Very largeshortfin makos over 3 m long have somewhat broader, more flattened and triangular upper teeth, perhaps more suitable forhandling small cetaceans than the awl-shaped teeth of smaller makos. Pinnipeds have not been recorded as mako food sofar (although they could be eaten in places where their distribution overlaps that of large makos), and makos have not beenseen scavenging on dead whales as with white sharks or tiger sharks.From prey types and other information, makos apparently feed at or near the surface and well below it, and may feed onbottom prey when close inshore. Stillwell and Kohler (1982) estimated that a 68 kg mako might consume about 2 kg of preyper day, and could eat about 8 to 11 times its body weight per year. Stillwell (1990) suggested that makos might consume upto 15 times their weight per year.This is a bold and responsive shark when a feeding stimulus (such as speared fish or a bait station) is present, andindividuals have charged divers repeatedly at high speed (usually veering off at the last moment) as well as chasing off lessdominant (possibly smaller?) conspecifics from baits. About a fourth of some 156 makos examined from the shark nets offKwaZulu-Natal, South Africa, had injuries, including missing fin tips, clasper tips, and mako bite patterns on the belly, flanks,pectoral fins, and gill regions. Some injuries were due to scavenging by other sharks, but mako-induced tooth cuts are likely

Sharks of the World, Vol. 2 113to be both from combat and mating, as with similar cuts on white and sand tiger sharks, including pectoral fin injuries thatmay be due to a male holding onto a female with its mouth during courtship. Open-mouth gaping at divers by these sharksmay be a threat display as in the white shark. Another possible threat display under non-baited conditions may be porpoisingor low short leaps at the surface, followed by figure 8 looping at speed. Behaviour is otherwise sketchily known, but it issuspected that a variety of behavioural elements, including varied social interactions, may be present in this species as inthe white shark. Shark netting data off South Africa suggests that makos were generally caught singly off beaches, thoughtwo occurred in the same net on the same day and a small group of five was caught in adjacent nets over nine days.The shortfin mako apparently has relatively few natural predators, perhaps because of its high activity level, speed and largesize. Their chief predators are humans, in the form of commercial fisheries. It is not immune from predation by its largercousin the white shark, which is very fast when chasing prey or conspecifics. Off California and in the Mediterranean Seasmall (less than 2 m long) makos have been found in white shark stomachs. In South Africa the fresh head of a mako (cut offat the pectoral fins) was found on a beach with unmistakable white shark wounds indicating a predation bout and noevidence of scavenging from fishing gear such as hook or net injuries. A live South African mako was caught with injuriessuggesting that a white shark grabbed it by the caudal fin and either lost it or let it go. Sailfish and swordfish may stab makoswith their beaks, presumably in self-defence; one adult female South African mako was found with a beak of a small sailfishsticking out of one eye. Makos caught in the KwaZulu-Natal shark nets were mostly dead (only 6% still alive whenexamined), vs. 16% alive for white sharks and 33% for spotted ragged-tooth sharks (Carcharias taurus). This probablyreflects different activity levels and oxygen requirements, with the mako (presumably with obligate ram-ventilation of its gills)having the highest levels and the more sluggish ragged-tooth (which can hover motionless) the least.Size: Maximum about 396 cm, estimated maximum total length about 408 cm. Size at birth between 60 and 70 cm, withfree-living individuals as small as 71 cm. Males adolescent at about 140 to 199 cm and maturing between 203 and 215 cmand reaching about 296 cm; females adolescent at about 233 to 286 cm, maturing at about 275 to 293 cm and reaching atleast 394 cm. Females apparently mature at smaller sizes in the Southern Hemisphere than in the western North Atlantic(Mollet et al., 2000). Several length-weight equations exist for this species and include:Stevens (1983): W(kg) = 4.832 x 10 -6 x TL(cm) 3.10Guitart (1975): W(kg) = 1.193 x 10 -6 x TL(cm) 3.46(n = 80, TL = 58 to 343 cm, Australia)(n = 23, TL = 160 to 260 cm, Cuba)Strasburg (1958): log W(lb) = -4.608 + 2.925 x log TL(cm)(Central Pacific)Cliff, Dudley and Davis (1990): W(kg) = 1.47 x 10 -5 x PCL(cm) 2.98(n = 143, PCL = 84 to 260 cm, South Africa)Kohler, Casey and Turner (1995): W(kg) = 5.2432 x 10 -6 x FL(cm) 3.1407where FL(cm) = 0.9286 x TL(cm) -1.7101 (n = 199)(n = 2081, western North Atlantic)Mollet et al. (2000): W(kg) = 7.2999 x TL (m) 3.224(n = 63, for females TL = 2.0 to 3.7 m, western North Atlantic)and W(kg) = 6.824 x TL(m) 3.137 (n = 64, for females TL = 2.0 to 3.4 m, Southern Hemisphere).Interest to Fisheries and Human Impact: The shortfin mako is an important species for pelagic longline fisheries whereverit occurs, because of its relative abundance and high quality meat, but is also caught in fisheries using drifting and set gillnets, and is taken by hook-and-line. It is taken as a utilized bycatch of tuna and swordfish longline fisheries worldwide, withcarcasses as well as fins being retained for market. The meat is utilized fresh, frozen, smoked and dried-salted for humanconsumption; the oil is extracted for vitamins; the fins used for shark-fin soup; the hides are processed into leather and thejaws and teeth are used for ornaments.Considerable fisheries for shortfin mako exist or existed in the Mediterranean Sea, off Cuba, in the Gulf of Mexico andCaribbean, off southern California, and in the western and central Pacific. It has been estimated that the Spanish longlinefleet caught about 750 t/y of shortfin makos in the Atlantic Ocean and Mediterranean Sea in the early 1990s (Bonfil, 1994).Off California, early juvenile shortfin makos were targeted by a short-lived experimental drift longline fishery and are a verywelcomed bycatch in the driftnet fishery for swordfish (Cailliet et al., 1993). Up to 475 t of shortfin makos were taken jointly bythese fisheries in 1987, and although CPUE did not show a declining trend concerns over the heavy exploitation of immaturefish prompted the closure of the experimental longline fishery in 1992 (O’Brien and Sunada, 1994). The total bycatch ofshortfin makos in the former high-seas driftnet fisheries in the North Pacific in the early 1990s was estimated at about 360 t/y(Bonfil, 1994). This species is apparently very common in the tuna fisheries of Indonesia: unconfirmed reports indicate thatlandings of shortfin makos from Indonesian waters attained 5 200 t in 1995 and that the estimated potential is about16 000 t/y (Priyono, 1998). The Brazilian longlining fleet based in Santos landed between 13.3 and 138.3 t of shortfin makosannually between 1971 and 1990 (Costa et al., 1996). Despite increasing fishing effort during this period, the CPUE ofshortfin makos has remained relatively stable with an initial slight decreasing trend followed by a slight increasing trend.Despite the role of shortfin makos in worldwide pelagic fisheries, catches are currently poorly reported to FAO at least.Brazil, New Zealand and the United States reported very small catches (2 to 76 t) to FAO from 1987 to 1997 (FAO FishStatPlus database, 2000).

114 FAO Species Catalogue for Fishery Purposes No. 1Big-game sports angling for mako sharks is widespread, with New Zealand and South Africa being traditional places foroffshore sports fishing for makos with heavy gear. The International Game Fish Association lists the shortfin mako as arecord game fish. In the 1980s mako angling became popular in the USA off southern California, with numerous anglersinvolved and mako tournaments rivalling competitive angling for marlin.Makos rarely bite divers and swimmers and few biting incidents are reliably reported. This suggests that the mako is notnormally inclined to bite people without an exciting stimulus or provocation and does not normally eat marine mammalianprey (except perhaps large females preying on small pelagic dolphins), however it should be treated with respect. Themostly offshore habitat of this species probably prevents it from coming in contact with recreational swimmers except underexceptional circumstances such as places where makos visit the vicinity of beaches with a narrow continental shelf. Itsspeed, power, large teeth, and boldness when a feeding stimulus is present should be cause for divers, especiallyspearfishers, to treat the mako with caution and observe what it is doing if it approaches. Divers should not dispute it forpossession of a shot fish and should back off if it is directing gapes their way, swimming in figure-8s, or giving bluff-charges.The shortfin mako tends to respond vigorously when hooked or harassed, and it should not be speared or otherwiseprovoked underwater. A defensive response by this animate torpedo when provoked may be far too quick for directedanti-shark weapons such as powerheads or repellent squirters to be effective. Steel-mesh armour suits were penetrated bythe powerful jaws and teeth of these sharks in tests. Electronic shark repellent devices worn by divers may be effective inthwarting oral contact by these sharks but may not be needed if the shark’s activities are correctly gauged.Makos occasionally bite boats, and mako boat incidents are second in number only to those involving the white shark. Mostof these incidents have occurred while makos were being played by anglers and should be regarded as human-provokedand abnormal. The angling and popular literature is rife with ‘mako stories’, in which these sharks bite, jump into, or evensmash right through the boats of their assailants. Anglers who suddenly find themselves sharing a boat with an aroused andvigorous mako have been known to leap into the water!Makos have become the subject of ecotouristic diving in recent years, with most of the mako dive sites being off southern Californiafrom the Los Angeles Basin to San Diego, but with sites also in South Africa and the Maldives. The sight of this sleek, beautiful,graceful shark underwater may be one of the high points for the shark-watching diver. The writer does not know offhand of anyattempts to display shortfin makos in public oceanaria, but small makos have been kept for physiological research in hydrodynamictunnels at Scripps Institution of Oceanography in southern California (United States).The conservation status of the shortfin mako is of moderate concern despite its relative abundance and vast distribution. Ithas been preliminarily assessed as a Lower Risk/Near Threatened species by the IUCN Shark Specialist Group. Theshortfin mako was found to have a midrange intrinsic rebound potential (a measure of its ability to recover from exploitation)among 26 shark species analysed (Smith, Au and Show, 1998). Fisheries are regulated and limited in the United States andNew Zealand, but not elsewhere where intensive epipelagic longline fisheries can have a major impact on it. Obviouslyinternational and regional management is needed to maintain sustainable catches. It is not known what the effects of theformer pelagic gill net fishery for scombroids was on makos and other oceanic sharks. Declines in landings have beenrecorded off the United States (east and west coasts) and in the eastern Atlantic by the Azorean ‘marrajo’ fleet, but trendsare little known elsewhere. Castro, Woodley and Brudek (1999) suggest that although it is very difficult to assess theconservation status of this shark because it is caught in numerous fisheries worldwide, it is reasonable to assume thatdecreases are occurring in those areas for which there is limited data or none whatsoever.Local Names: Mako (original Maori name for this species, New Zealand); Shortfinned mako shark, Mako shark, Mako,Atlantic mako shark, Pacific mako shark, Pacific bonito shark, Pacific mako, Mackerel shark, Blue shark, Sharp nosed,Sharpnosed or Sharp-nosed mackerel shark, Mackerel porbeagle, Mediterranean mackerel shark (English); Ossirina dellospallanzani, Meanto, Pisci tunnu, Piscicani, Cani di mari, Cane di mare di Messina, Cagnia (Italy); Psina cavlozuba, Psinadugonoska (Adriatic); Anequin (Portugal); Marracho, Rinquim, Mako, Shortfin mako, Marrajo criollo (Azores); Sobraig,Sobratg, Ludia marraco, Marraquet, Marrajo, Tiburón (Spain); Alecrín (Mexico); Dentudo, Dientuso azul (Cuba); Ganumusorrah (India, Tamil); Blue pointer, Mako shark, Snapper shark (Australia); Porpoise shark, Blue porpoise shark, Sharpnosemako, Mambone, Moro (South Africa); Aozame, Morozame, Awozame (Japan); Bonito shark (California); Anequinbarbatana curta (Mozambique).Remarks: Moreno and Morón (1992b) give detailed descriptions of the coloration of typical I. oxyrinchus and the Azores‘marrajo criollo’ mako, which was long recognized by Azores fishermen as a separate form as distinguished from typicalI. oxyrinchus or marrajo, and marrajo negro or I. paucus. Apparently Cuban and Japanese fishermen recognized thelongfin and shortfin makos as being distinct long before scientists were aware of their existence.Literature: Garman (1913); Barnard (1925); Fowler (1936, 1941); Whitley (1940); Bigelow and Schroeder (1948); Smith(1949, 1957d); Strasburg (1958); Farquhar (1963); Stead (1963); Garrick and Schultz (1963); Randall (1963); Applegate(1966, 1977); Guitart (1966, 1975); Garrick (1967); Carey and Teal (1969); Lineaweaver and Backus (1970); Mundus andWisner (1971); Carey et al. (1972); Gubanov (1974, 1978); Bass, D’Aubrey and Kistnasamy (1975a); Ellis (1976); Penrith(1978); Glikman and Dolganov (1980); Cadenat and Blache (1981); Stillwell and Kohler (1982); Cadenat and Blache (1981);Carey (1982, 1990); Pratt and Casey (1983); Gilmore (1983); Stevens (1983, 1984, 1990, 1992); Cailliet and Bedford(1983); Compagno (1984, 1990a, b); Nakaya (1984); Block and Carey (1985); Compagno, Ebert and Smale (1989); Quero(1984); Paulin et al. (1989); Springer (1990); Fischer et al. (1990); Cliff, Dudley and Davis (1990); Stillwell (1990);Compagno, Ebert and Cowley (1991); Casey and Kohler (1992); Moreno and Morón (1992b); Holts and Bedford (1992);Anderson and Ahmed (1993); Cailliet, Holts and Bedford (1993); Francis and Randall (1993); Michael (1993); Bonfil (1994);

Sharks of the World, Vol. 2 115Last and Stevens (1994); Seret (1994); Costa et al. (1996); Priyono (1998); Smith, Au and Show (1998); Castro, Woodleyand Brudek (1999); Mollet et al. (2000); H. Mollet (pers. comm.).Isurus paucus Guitart, 1966 Fig. 75Isurus paucus Guitart, 1966, Poeyana, Ser. A, (15): 3, figs. 1, 2A, 3A, 3C. Syntypes: 2 260 mm PCL adult female, 2 030 mmPCL adult male, and 1 955 mm PCL adult female, possibly in the Instituto de Biología or Instituto de Oceanología, Cuba,collected in the Caribbean near Cuba. No types known according to Eschmeyer (1998: CD-ROM).Synonyms: ?Lamiostoma belyaevi Glikman, 1964: 105. Syntypes: a set of fossil tooth crowns from the bottom of theWest-Central Pacific at 5 120 m depth, 13°00’ N, 176°04’ E. Syntypes doubtfully referred to the living species (see remarksbelow). Isurus alatus Garrick, 1967: 677. Holotype: US National Museum of Natural History, USNM-197427, 1 233 mm TLimmature male, Indo-Pacific, no precise locality. Status confirmed by Howe and Springer (1993: 3).Other Combinations: None.FAO Names: En - Longfin mako; Fr - Petit taupe; Sp - Marrajo carite.UPPER AND LOWER TEETH ON LEFT SIDEUNDERSIDE OF HEADFig. 75 Isurus paucusField Marks: Slender, spindle-shaped body, moderately long conical snout, large blade-like teeth without lateral cusplets orserrations, long gill slits, pectoral fins broad-tipped and as long or longer than head, large first dorsal fin with light free reartip, minute, pivoting second dorsal and anal fins, strong keels on caudal peduncle, short secondary keels on caudal base,crescentic caudal fin, ventral surface of body dusky on underside of head.Diagnostic Features: Snout broadly pointed. Eyes relatively large. Mouth parabolic in shape. Lower anterior teeth slightlyprotruding from jaws and in line with the laterals; anterior teeth with relatively broad, nearly straight cusps with unflexed tips;cusps of first upper anterior teeth with complete cutting edges; intermediate teeth with nearly straight or slightly hookedcusps. Body slender and elongated. Pectoral fins nearly straight and very broad-tipped, anterior margins about 23 to 31% oftotal length and equal to or greater than head length. Origin of first dorsal fin well behind the pectoral free rear tip; firstdorsal-fin apex broadly rounded and hardly angular at all stages; first dorsal-fin height greater than base length at all stages

116 FAO Species Catalogue for Fishery Purposes No. 1(smaller in term foetuses). Vertebral total count 195 to 197. Colour: dorsolateral coloration dark slaty blue or grey-black inlife, underside white but with underside of snout and jaws dark in adults and large juveniles though not in young; dark colourof head entirely covering gill septa; dark colour of flanks extending ventrally onto abdomen in adults; pelvic fin completelydark, underside white with prominent dark margin; first dorsal fin as dark as back; anal fin dark except for white free rear tipand posterior margin.Distribution: Oceanic and tropical, probably circumtropical but records sporadic and distribution sketchily known, probablyoften mistaken for the apparently far more common Isurus oxyrinchus or included with records for it. Western Atlantic:Florida, Gulf Stream off eastern USA, Cuba, southern Brazil. Eastern Atlantic: Spain, Portugal, probably Mediterranean,Morocco, Western Sahara, Canary Islands, Mauritania, Guinea-Bissau, Liberia, Ghana, ?Cape Verde Islands. WesternIndian Ocean: ? South Africa, Madagascar. Western Pacific: Japan, Taiwan (Province of China), Australia (Queensland andnorthern New South Wales, also possibly off northern Australia). Central Pacific: Northeast of Micronesia, between Solomonand Nauru Islands, area south of Johnston and Hawaiian Islands, near Phoenix Island, and north of Hawaiian Islands.Eastern Pacific: United States (southern California).Habitat: A little-known epipelagic, tropical and warm-temperate shark, apparently common in the western Atlantic andpossibly in the Central Pacific, but rare elsewhere. Said to be deep-dwelling but bathymetric data was not available.Biology: The biology of the longfin mako is poorly known. In the eastern Atlantic this species is possibly rare compared to I.oxyrinchus, and landings of longfin mako in Spanish fishing ports sampled by Moreno and Morón (1992) included only 51specimens compared with 45 679 shortfin mako (0.1%). The often slimmer build and broad, long pectoral fins of this sharksuggest that it is slower and less active than its better-known relative, the shortfin mako (J. Casey, pers. comm.). Itsmacroceanic morphology suggests similar slow cruising in the epipelagic zone as in the oceanic whitetip (Carcharhinuslongimanus) and the blue shark (Prionace glauca) rather than the more active, scombroid-like swimming of Isurusoxyrinchus. The longfin mako is apparently endothermic, with countercurrent vascular heat exchangers for its bodymusculature, eyes, brain and viscera as in other lamnids (Carey, 1982), but the levels of temperature elevation it canachieve above ambient conditions have apparently not been measured.The longfin mako is ovoviviparous, with uterine cannibalism; foetuses are larger than those of I. oxyrinchus, are full-term at92 to 120 cm, and occur as a litter of 2 to 8 young. It may approach land to give birth.Food of this shark is presumably schooling fish and pelagic cephalopods. Michael (1993) noted that one was found with aswordfish sword stuck in its abdomen, though it is not known if swordfish are an important item of this mako’s diet as with theshortfin mako.Size: Maximum reported 417 cm; size at birth between 97 and 120 cm; a male (Central Pacific) was adult at 245 cm; adultfemales (western North Atlantic) were 245 to 417 cm long.Interest to Fisheries and Human Impact: Probably taken regularly in tropical pelagic longline fisheries for tuna andswordfish as bycatch (with some marketed in Tokyo). Historically it was often taken in the Cuban longline fishery for sharksoff the north coast of Cuba and averaged about a sixth of the total weight of sharks caught there in 1971-1972. Whether it isstill as common there at present is unknown. It is utilized fresh, frozen and dried-salted for human consumption but the meatis of lower quality than the shortfin mako and it is often finned and discarded at sea. In addition to longlines, the species istaken with hook-and-line and with anchored gill nets.

Sharks of the World, Vol. 2 117This species has not bitten people or boats and has not, to the writer’s knowledge, been observed underwater or kept incaptivity.The conservation status of this species is uncertain but needs urgent investigation. It has probably been captured asbycatch in oceanic fisheries wherever it occurs and is a minor bycatch of fisheries targeting shortfin makos in the easternAtlantic. No trends are obvious for the species as there are no catch statistics for it except off Cuba and the United Statesand these are not long-term. The United States reported small catches (2 to 12 t in 1987 to 1997) to FAO (FAO FishStat Plusdatabase, 2000). Problems for the longfin mako include the rarity or uncommonness of this shark in most places (exceptperhaps the western Atlantic in the 1970s at least); limited knowledge of its biology; large maximum size; apparently lowerfecundity than the shortfin mako; and occurrence as a limited but complementary bycatch of high-intensity oceanic fisheriestargeting shortfin makos, other sharks, and pelagic scombroid fishes. These problems should arouse the concern ofconservation and fisheries organizations, because of increased mortality of an uncommon or rare species due to finning andpossibly capture trauma.Local Names: Longfinned mako shark, Taupe longue aile (France); Dientuso prieto (Cuba); Marrajo negro (Azores);Bake-aozame (Japan).Remarks: Garrick (1967), Compagno and Vergara (1978), and Compagno (1981a) thought that the species Lamiostomabelyaevi Glikman, 1964 might prove to be an earlier name for I. paucus, particularly because a stuffed Isurus illustrated in aphotograph in Glikman (1964, figs 31-32) and labelled L. belyaevi appeared to be a longfin mako. This may be irrelevanteven if correct. A translation of Glikman’s description of L. belyaevi (pp. 105, 132-133) by Mrs L.J. Dempster with the aid ofDr V.V. Barsukov (noted in Compagno, 1984) revealed that Glikman deliberately refrained from naming the stuffed Isurus asholotype of L. belyaevi but instead picked one lot of teeth crowns dredged from the ocean bottom 5 120 m deep at RVVITYAZ station 5128, 13°00’ N, 176°04’E in the Central Pacific southwest of the Hawaiian Islands (Glikman, 1964, pl. 31,figs 13, 14, 18, 19) for this role. Examination of Glikman’s photos did not convince Compagno (1984) that the shark or sharksrepresented by these teeth were necessarily conspecific with I. paucus and were not conspecific with I. oxyrinchus or evensome extinct Isurus species. Hence the substitution of the species name belyaevi for paucus was rejected, especiallybecause the former is based on such poor material. It is uncertain if the stuffed specimen illustrated by Glikman is I. paucusalso, because some of the characters ascribed to it (snout very long and acute, pectoral fins falcate, and pectoral fin lengthslightly less than the distance from snout tip to upper margin of first gill opening, vs. snout short and bluntly conical, pectoralfins not strongly falcate, and pectoral-fin length much longer than the distance from snout tip to upper margin of first gillopening in I. paucus) indicate that it might be a specimen of I. oxyrinchus instead.Literature: Glikman (1964); Guitart (1966, 1968, 1975); Garrick (1967); Bass, D’Aubrey and Kistnasamy (1975a);Fourmanoir and Laboute (1978); Dodrill and Gilmore (1979); Gilmore (1983, 1993); Pinchuk (1983); Compagno (1984);Nakaya (1984); Quero (1984); Sadowsky, Amorim and Arfelli (1986); Killam and Parsons (1986); Joung, Wu and Chen(1989); Springer (1990); Moreno and Morón (1992); Michael (1993); Seret (1994); Stevens and Scott (1995); Stevens andWayte (1998); Castro, Woodley and Brudek. (1999); Mollet et al. (2000); R. Bonfil (pers. comm.); H. Mollet (pers. comm.); D.Ebert (pers. comm. on California record).Lamna Cuvier, 1816Genus: Subgenus Lamna Cuvier, 1816 (Genus Squalus Linnaeus, 1758), Reg. Anim., ed. 1, 2: 126.Type Species: Squalus cornubicus Bloch and Schneider, 1801 by monotypy, equals S. cornubicus Gmelin, 1788: 1497,and a junior synonym of S. nasus Bonnaterre, 1788.Number of Recognized Species: 2.Synonyms: Genus Lamia Risso, 1827: 123. Type species, “L. cornubicus L” = Squalus cornubicus Gmelin, 1788 bymonotypy. A junior homonym of Lamia Fabricius, 1775 in Coeloptera (Insecta). Genus Selanonius Fleming, 1828: 169.Type species, Selanonius walkeri Fleming, 1828 by monotypy (Eschmeyer, 1998: CD-ROM), junior synonym of Squalusnasus Bonnaterre, 1788. Genus Exoles Gistel, 1848: ix. Replacement name for Lamia Risso, 1827, and hence taking thesame type species, Squalus cornubicus Gmelin, 1788.Diagnostic Features: Snout very bluntly conical to acutely conical. Eyes large, 1.6 to 2.8% of total length. Nostrils situatedmedial to horizontal head rim in ventral view. Mouth width 1.4 to 2.3 times its length. Anterior teeth not enlarged; anterior andintermediate teeth not compressed but upper lateral teeth somewhat compressed, teeth not forming a continuous cuttingedge; intermediate teeth small and about half height of adjacent anteriors, intermediate teeth with normal cusps directedposterodistally or vertically; second lower anterior teeth moderately enlarged and taller than second upper anterior tooth.Total tooth count 49 to 60; roots of anterior teeth deeply arched, with narrow short lobes; lateral cusplets present on teeth atall stages except newborn sharks; teeth smooth-edged; cusps of anterior teeth not strongly flexed. Body very stout. Firstdorsal-fin origin over or just behind the pectoral-fin insertions. Anal-fin origin about under second dorsal-fin origin.

118 FAO Species Catalogue for Fishery Purposes No. 1Secondary caudal keels present and strong. Total vertebral count 150 to 173. Cranium with rostral cartilages more or lessexpanded and hypercalcified. Intestinal valve of ring type with count 38 to 41. Length of adults 1.5 to at least 3.1 m. No blackaxillary spot at pectoral-fin insertions; pectoral-fin tips not abruptly black on their ventral surfaces but sometimes with darkmargins.Local Names: Porbeagles, Mackerel sharks, Salmon sharks.Remarks: As with the makos (Isurus) there has been confusion in the past as to the number of valid species of mackerelsharks (Lamna). Material examined by the writer and the accounts of Bigelow and Schroeder (1948), Nakaya (1971),Stevens, Dunning and Machira (1983), and Compagno (1984, 1990b, c), suggest that there are only two species, L. ditropisand L. nasus, with L. philippi from Chile and L. whitleyi from New Zealand and Australia most probably being synonyms ofL. nasus. Lamna ditropis was compared in detail to L. nasus by Nakaya (1971) and the two are readily separable inexternal morphology and coloration. Nakaya (1971) also noted differences between the dentitions of the two species, withL. ditropis having more oblique-cusped upper first lateral teeth. This needs to be confirmed by a comprehensive study of thedentition of Lamna from different areas, as some Southern Hemisphere L. nasus have upper first lateral teeth about asoblique-cusped as L. ditropis. Compagno (1984 and this volume) noted differences in the colour of the first dorsal rear tipand the flanks between the two species (see below), while Compagno (1990b) described and illustrated striking differencesin their cranial morphology.Pillai and Honma (1978) reported L. ditropis from the southern Indian Ocean without data confirming their identification.Compagno (1984) suggested that the species in question was L. nasus, and had examined a specimen from the southernIndian Ocean (open sea between Kerguelen and St. Paul Islands, ANTON BRUUN Cruise 5, Sta. 309, 960 mm immaturefemale, 42°23’S, 74°56’E, in collections of the California Academy of Sciences). There are also records from near KerguelenIsland (Duhamel and Ozouf-Costaz, 1982). Records of L. nasus from sub-Antarctic seas were summarized by Duhamel andCompagno (1985) and Compagno (1990c).A few specimens of large L. nasus (1.6 to 2.1 m long) have subsequently been examined by the writer from the SouthernHemisphere, off the Western Cape, South Africa, off Marion Island, and between the Prince Edward and Crozet Islands.These specimens agreed with L. nasus from the North Atlantic in having more elongated snouts, moderately hypercalcifiedrostra, no light flank markings, and a light first dorsal rear tip, but agree with L. ditropis in having dusky or blackish faces withdark coloration on the underside of the head and a spotted, dusky abdomen. Stevens, Dunning and Machira (1983)illustrated a similar colour pattern in a porbeagle from the Tasman Sea. These ‘black-faced’ porbeagles resemble the‘marrajo criollo’ makos from the Azores (Isurus oxyrinchus), which have a colour pattern similar to I. paucus (see above). Itmay be that black-faced Lamna nasus have been misidentified as L. ditropis in the Southern Hemisphere, but it is notknown if large southern L. nasus are exclusively dark-faced, or if this dark form is geographically restricted within the vastsouthern range of the porbeagle. Apparently it is necessary to conduct comparative studies of porbeagles from differentareas, including molecular research, to sort out these problems. One aspect of this is to determine if separate populations orsubpopulations exist for L. nasus, within contiguous ranges and north or south of the Equator.It is an interesting irony of nomenclature that the names Lamna Cuvier, 1816 and its synonym Lamia Risso, 1827, based onthe classical Lamia (Carcharodon carcharias) and connoting a frightful anthropophagous monster, should be used for therather less assertive porbeagles and salmon sharks.snout longKey to Species:1a. Snout relatively long, distance from snout tipto eye 50% or more of the distance from eye tofirst gill opening (Fig. 76); free rear tip of firstdorsal fin abruptly white; sides abovepectoral-fin bases dark, without an extensionof the white abdominal area . . . . . . . . . . . . . . . . . Lamna nasusFig. 76 Lamna nasus1b. Snout shorter, distance from snout tip to eye40% or less of the distance from eye to first gillopening (Fig. 77); free rear tip of first dorsal findark; sides above pectoral-fin bases markedwith a white patch extending forward from theabdominal area . . . . . . . . . . . . . . . . . . . . . . . Lamna ditropissnout shortFig. 77 Lamna ditropis

Sharks of the World, Vol. 2 119Lamna ditropis Hubbs and Follett, 1947 Fig. 78Lamna ditropis Hubbs and Follett, 1947, Copeia, 1947(3): 194. Holotype, Museum of Comparative Zoology, HarvardUniversity, MCZ-36471, adult male (partial specimen, size uncertain); type locality, La Jolla, California, 92 to 107 m off the LaJolla Beach Club in shallow water.Synonyms: None.Other Combinations: None.FAO Names: En - Salmon shark; Fr - Requin-taupe saumon; Sp - Marrajo salmón.UPPER AND LOWER TEETH ON RIGHT SIDE, LINGUAL VIEWUNDERSIDE OF HEADField Marks: Heavy spindle-shaped body, short conical snout, moderately large blade-like teeth with lateral cusplets, longgill slits, large first dorsal fin with dark free rear tip, minute, pivoting second dorsal and anal fins, strong keels on caudalpeduncle, short secondary keels on caudal base, crescentic caudal fin, underside of preoral snout dark, often duskyblotches on ventral surface of body and white patches over pectoral bases.Diagnostic Features: Snout short and bluntly pointed, with preoral length 4.5 to 7.6% of total length (adults 4.5 to 5.0%),space from eye to first gill slit 1.3 to 1.9 times preorbital length. First upper lateral teeth with oblique cusps. Total vertebralcount 170, precaudal vertebral count 103. Cranial rostrum expanded as a huge hypercalcified knob which engulfs most ofthe rostral cartilages except bases in adults. Colour: dark grey or blackish on dorsolateral surface of body, white below, withwhite abdominal colour extending anteriorly overpectoral bases as a broad wedge-shaped band; firstdorsal fin without a white free rear tip; ventral surface ofhead dusky and abdomen with dusky blotches in adultsbut not in young.Distribution: Coastal and oceanic. North Pacific: Japan(including Sea of Japan), North Korea, South Korea, andthe Pacific coast of Russia (including Sea of Okhotsk) toBering Sea and the eastern Pacific coast of the USA andCanada (Alaska south to British Columbia, Washington,Oregon, and southern California) and probably Mexico(northern Baja California).Fig. 78 Lamna ditropis

120 FAO Species Catalogue for Fishery Purposes No. 1Habitat: A common coastal-littoral, offshore and epipelagic shark with a preference for boreal to cool temperate waters,found at depths from the surface to below 152 m. One was photographed at 255 m near the bottom in Monterey Canyonusing an underwater camera, while a diver in a submersible saw one at 224 m off Alaska. Salmon sharks are common incontinental offshore waters but range inshore to just off beaches; they also are abundant far from land in the North PacificOcean basin, along with their pelagic fish prey.Biology: Salmon sharks are common and are often encountered by oceanic and coastal fisheries but are sketchily knownbiologically. Behaviour and sociobiology are little-known. They occur singly and in schools or feeding aggregations ofseveral individuals and in some areas are seen at or near the surface. Water temperatures where salmon sharks werecaught ranged from 2.5° to 24°C. They are swift-swimming sharks, maintaining a body temperature well above ambientwater temperature. Recent studies suggest that salmon sharks may have the highest body temperature of any shark. Bodytemperature elevations of 8° to 11°C above that of the surrounding water have been reported for smaller specimens, whileelevations up to 13.6°C have been recorded in larger ones (Smith and Rhodes 1983; Goldman and Human, in press).Salmon sharks are migratory, with segregation by size and sex, and with larger sharks ranging more northerly than young. Inthe western North Pacific large sharks migrate from Japanese waters (where they breed) in the wintertime, move north to theSea of Okhotsk and the western Bering Sea when the water warms, and return to Japan in the autumn or early winter (for aone-way distance of 3 220 km). In the eastern Pacific females apparently migrate south to pup in the spring off Oregon andCalifornia, USA, as suggested by commercial fish catch records, washed up (beached) young of the year and anecdotalinformation. A strong sexual segregation appears to exist across the Pacific Ocean basin, with males dominating thewestern North Pacific and females dominating the eastern North Pacific (K.J. Goldman and J. A. Musick, pers. comm.).This shark reproduces by aplacental viviparity, with uterine cannibalism (oophagy); litter size is 2 to 5 young. Length ofgestation period might be nine months; length of entire reproductive cycle unknown. Breeding occurs in late summer andinto autumn, and females bear young in spring. Breeding and nursery areas may be localized in the offshore western NorthPacific between about 156° and 180°E in the open ocean, off the southern Kuril region, and in the Sea of Okhotsk, whereyoung below 60 cm (possibly newborn) occur and juveniles up to 110 to 120 cm long also are found. Age 0 and 1 salmonsharks occur off California, USA, suggesting that a breeding and nursery ground might exist in the eastern North Pacific (K.J.Goldman and J.A. Musick, pers. comm.). Males may mature at 5 years and about 182 cm TL, and females at 8 to 10 yearsand about 221 cm TL (Tanaka, 1980). Females in the eastern North Pacific live to at least 20 years of age, males to at least27 years; preliminary indications are that female salmon sharks mature at an earlier age and are heavier in the eastern NorthPacific relative to those living in the western North Pacific (K.J. Goldman and J.A. Musick, pers. comm.).Salmon sharks are opportunistic feeders and eat a variety of pelagic and demersal bony fishes including Pacific salmon andsteelhead trout (Salmonidae), herring and sardines (Clupeidae), pollock, Alaska cod and tomcod (Gadidae), lancetfishes(Alepisauridae), daggerteeth (Anotopteridae), sauries (Scombresocidae), lanternfishes (Myctophidae), pomfrets(Stromateidae), mackerel (Scomber, Scombridae), lumpfishes (Cyclopteridae), sculpins (Cottidae), possibly rockfish(Sebastes, Scorpaenidae), possibly sablefish (Anoplopomatidae), and Atka mackerel (Pleurogrammus, Hexagrammidae).Salmon sharks also feed on spiny dogfish (Squalus acanthias, Squalidae) and several species of pelagic squid, and havebeen attracted to bycatch offal dumped by shrimp trawlers.The salmon shark is generally considered to be one of the principal predators of Pacific salmon (Oncorhynchus) apart fromhumans and is depicted as voraciously feeding on salmon. This is apparently the case around the Aleutians and the Gulf ofAlaska, where peaks in abundance in salmon sharks follow maximum catches of salmon and the distribution and migrationsof the two appear to be strongly correlated as predator and prey. Salmon sharks caught by Japanese pelagic salmon gillnetters in this area have had salmon in their stomachs and little else. However Blagoderov (1994) suggested that thisrelationship is highly unlikely, and cited major differences in areal distribution between salmon and salmon sharks in thewestern North Pacific, with most salmon sharks concentrated south of the main migration path of salmon and very few withinit. In the western North Pacific these sharks congregate in areas with breeding aggregations of herring and sardines andmay be selecting these fishes rather than salmon.Size: Maximum total length about 305 cm; anecdotal accounts mention sizes of 3.7 to 4.3 m TL but cannot be confirmed,and confusion with the larger white shark is possible and has happened. Size at birth between 40 and 50 cm and 85 cm TL,with the largest foetuses at least 70 cm long and the smallest free-ranging young between 40 and 50 cm. Males maturing atabout 182 cm TL (5 years) and females at about 221 cm TL (8 to 10 years); both sexes adult over about 210 to 220 cm TL.Interest to Fisheries and Human Impact: This species has been fished in the North Pacific in the past by Japanese coastaland oceanic longliners. Salmon sharks are commonly caught by Japanese, United States and Canadian offshore salmon gillnetters as bycatch but are generally discarded (except for fins). They are also caught in salmon seines, by salmon trollerstowing hooks, and possibly by bottom trawlers off Alaska; Russian research vessels have regularly caught them in pelagictrawls in the western North Pacific. They are occasionally trammel-netted by halibut fishermen off California and haveshowed up in numbers as bycatch in gill nets set for swordfish and threshers off California but have usually not beenmarketed there. Sports anglers in Alaska and Canada catch salmon sharks using rod and reel much like porbeagle anglersin the North Atlantic.Salmon sharks are generally considered a nuisance for the damage they do to salmon nets and other fishing gear. Acommercial fishery was initiated off Alaska but this did not succeed. FAO catch statistics for recent salmon shark landingswere not available (FAO FishStat Plus database, 2000) but available data (Makihara, 1980) indicates that Japanese fisherslanded 100 to 41 000 t during 1952-1978 (with one very high catch year, 41 000 t in 1954, but mostly below 10 000 t and

Sharks of the World, Vol. 2 121averaging about 6 900 t). Bycatch of salmon sharks in the flying squid and large-mesh driftnet fisheries of the North Pacific in1990, just before high-seas driftnets were internationally banned was estimated to be about 5 400 t and 71 t respectively.The flesh of the salmon shark is used fresh for human consumption in Japan, where it is processed into various fishproducts, and to a lesser extent in Alaska and California, United States, where it is seldom marketed and has in the past(California) been occasionally sold as swordfish. Presumably its flesh is less desirable than that of the shortfin mako. Its oil,skin (for leather), and fins (for shark fin soup) are utilized also. The heart of the salmon shark is highly appreciated in a localsashimi dish in the northern fishing port of Kesennuma, where most of the landings of salmon sharks occur in Japan(R. Bonfil, pers. comm.).The salmon shark has been regarded as potentially injurious to people because of its large size and relationship to otherspecies that occasionally bite people, but has never been positively identified in shark-bite incidents. There are a fewunsubstantiated incidents reported for the species, but possibly by confusion with the white shark. Salmon sharks arereported as occasionally circling, approaching and bumping fishing vessels and sports boats off southeastern Alaska (Paustand Smith, 1985), but their identity needs to be confirmed to eliminate white sharks as being involved in such activities.Divers have seen and photographed schools of adult salmon sharks underwater, with no agonistic overtures on the part ofthe sharks (R. Lea, pers. comm.). An ecotouristic dive site at Roca Partida, Socorro Islands, Mexico has reported “Lamnanasus”, but it is not known if salmon sharks or other lamnids are being viewed there. The salmon shark is not currently held incaptivity in large oceanaria, nor does the writer know of any attempts to keep salmon sharks in the past.Recently (1997) there has been numerous strandings of small salmon sharks, ca. 1 m long, off north-central and southernCalifornia (R. Lea, pers. comm.), which was of rare occurrence in the 1970s and 1980s. Whether this has to do withhuman-induced environmental problems such as pollution or unusual water conditions is not known.The conservation status of this species is of concern because it is heavily fished as largely discarded but complementarybycatch (with finning) in major pelagic fisheries in the North Pacific. Unlike Lamna nasus, this species has limited fisheriesstatistics (with no country reporting catch statistics to FAO in 1997) and no regulation of the largely pelagic fishery ininternational waters, so that trends in abundance are unknown. It also has a negative image as an abundant and low-valuepest that avidly eats or damages valuable salmon and wrecks gear, which encourages fishers to kill it and add to mortalityfrom finning and capture trauma. Knowledge of its biology is limited despite its abundance, which invites neglect, but itsfecundity is very low and probably cannot sustain current fishing pressure for extended periods.The Alaska Board of Fisheries has closed all commercial shark fishing and has heavily regulated the sport fishery in Alaskastate waters since 1997 in view of the lack of biological information at a time when there was a small amount of commercialfishing and a large increased interest in sport fishing for salmon sharks (Goldman and Human, in press). The North PacificFishery Management Council is currently considering closure of commercial fishing for sharks in Federal waters as noFederal Management plan exists specifically for sharks in the Gulf of Alaska and the Aleutian Islands. Currently, salmonsharks are allowed as bycatch, and are included in the commercial bycatch TAC (Total Allowable Catch) for Alaska Federalwaters. Sport fishing regulations in Alaska include EEZ waters and are two sharks per person per year, one in possession atany time (one per day).Local Names: Nezumizame, Mokazame, Radukazame, Sakezame, Japanese mackerel shark (Japan); Salmon shark,Porbeagle, Mackerel shark (English).Literature: Hubbs and Follett (1947); Bigelow and Schroeder (1948); Roedel and Ripley (1950); Strasburg (1958); Lindbergand Legeza (1959); Sano (1962); Farquhar (1963); Larkins (1964); Kato, Springer and Wagner (1967); Nakaya (1971,1984); Miller and Lea (1972); Hart (1973); Makihara (1980); Tanaka (1980); Urquhart (1981); Smith and Rhodes (1983);Compagno (1984); Paust and Smith (1986); Paust (1987); Brodeur (1988); Applegate et al. (1989); Hanan, Holts and Coan(1993); Blagoderov (1994); Bonfil (1994); Nagasawa (1998); Castro, Woodley and Brudek (1999); Goldman and Human, (inpress); K.G. Goldman and J. A. Musick (pers. comm.); S. Kato (pers. comm.); R. Lea (pers. comm.); T. Neal (pers. comm.);H. Mollet, (pers. comm.).Lamna nasus (Bonnaterre, 1788) Fig. 79Squalus nasus Bonnaterre, 1788, Tabl. Encyclop. Method. Trois Reg. Nat., Ichthyol., Paris: 10, pl. 85, fig. 350. Holotypeunknown, type locality probably Cornwall, England (Eschmeyer, 1998: Cat. Fish.: CD-ROM).Synonyms: Squalus glaucus Gunnerus, 1768: 1, pl. 1. Type locality, Norway. A junior homonym of S. glaucus Linnaeus,1758 ( = Prionace glauca). Squalus cornubicus Gmelin, 1788: 1497. No types known according to Eschmeyer (1998:CD-ROM). Type locality: Cornwall, England. Squalus pennanti Walbaum, 1792: 517. Type locality: Atlantic. No typesaccording to Eschmeyer (1998: CD-ROM). Squalus monensis Shaw, 1804: 350. Based on the ‘Beaumaris Shark’ of

122 FAO Species Catalogue for Fishery Purposes No. 1Pennant, a 2.33 m (7 ft) shark observed and reported by the Rev. Hugh Davies, of Beaumaris, Isle of Anglesey, Wales. Notypes known according to Eschmeyer (1998: CD-ROM). Shaw thought that his S. monensis might be the same as Squaluscornubicus Gmelin, 1788 (= Lamna nasus), and that differences between them might be attributable to sexual dimorphism.Squalus cornubiensis Pennant, 1812: 152. Type locality: Cornwall. Variant spelling of S. cornubicus Gmelin, 1788according to Eschmeyer (1998: CD-ROM). Squalus selanonus Leach, 1818: 64, pl. 2, fig. 2. Holotype: University ofEdinburgh, ca. 2.6 m TL adult male, Lochfyne, Scotland (also, Eschmeyer, 1998: CD-ROM). Selanonius walkeri Fleming,1828: 169. Argyll, Scotland. Based on Squalus selanonus of Walker, 1769 (manuscript name) according to Eschmeyer(1998: CD-ROM). Lamna punctata Storer, 1839: 185, pl. 3, fig. 2. Also Storer, 1839: 534, pl. 8. New combination for andmisinterpretation of Squalus punctatus Mitchill, 1815 (= Carcharhinus isodon), itself a junior homonym of S. punctatusBloch and Schneider, 1801(= Ginglymostoma cirratum). Type locality: Massachusetts Bay; cf. Eschmeyer (1998:CD-ROM). Lamna pennanti Desvaux, 1851: 23. Possibly new combination based on Squalus pennanti Walbaum, 1792.Oxyrhina daekayi Gill, 1862a: 60. New name for Lamna punctata Storer, 1839 according to Eschmeyer (1998: CD-ROM).Isuropsis dekayi Gill, 1873: 813 (emended spelling of specific name). Lamna philippi Perez Canto, 1886: 1. Type locality,Chile. Types? Lamna whitleyi Phillipps, 1935: 239, fig. 3. Syntypes: Whereabouts unknown according to Eschmeyer (1998:CD-ROM). Type Locality, Island Bay, Wellington, New Zealand.Other Combinations: Lamna cornubica (Gmelin, 1788).FAO Names: En - Porbeagle; Fr - Requin-taupe commun; Sp - Marrajo sardinero.DERMALDENTICLESUPPER AND LOWER TEETH ON LEFT SIDENOSTRILUNDERSIDE OF HEADFig. 79 Lamna nasusField Marks: Heavy spindle-shaped body, moderately long conical snout, moderately large blade-like teeth with lateralcusplets, long gill slits, large first dorsal fin with abruptly white free rear tip, minute, pivoting second dorsal and anal fins,strong keels on caudal peduncle, short secondary keels on caudal base, crescentic caudal fin, ventral surface of body whiteand not extending over pectoral bases as white patches.Diagnostic Features: Snout long and sharply pointed, with preoral length 5.9 to 9.0% of total length (adults 5.9 to 7.3%) andspace from eye to first gill slit 1.7 to 2.5 times preorbital length. First upper lateral teeth with nearly straight cusps. Totalvertebral count 150 to 162, precaudal vertebral count 85 to 91. Cranial rostrum with enlarged but discrete hypercalcifiedrostral cartilages, not forming a massive knob. Colour: grey or bluish grey to blackish above, white below, with whiteabdominal colour terminating at rear end of pectoral bases; first dorsal fin with an abruptly white or greyish white free rear tip;ventral surface of head white and abdomen without dusky blotches in adults of typical Northern Hemisphere porbeagles, butunderside of head dark and abdomen blotched in some Southern Hemisphere adults.

Sharks of the World, Vol. 2 123Distribution: Coastal and oceanic, amphitemperate, with centres of distribution in the North Atlantic and in a circumglobalband of temperate water of the southern Atlantic, southern Indian Ocean, southern Pacific and Antarctic Ocean. WesternAtlantic: Greenland, Canada (Newfoundland Banks, Gulf of St. Lawrence and Nova Scotia), United States (Maine,Massachusetts, Rhode Island, rarely New York, New Jersey and possibly South Carolina), and Bermuda; southern Braziland Uruguay to southern Argentina. Eastern Atlantic: Iceland and western Barents Sea to Baltic and North Seas, EnglishChannel, Straits of Gibraltar, and Mediterranean Sea, including Russia, Norway, Sweden, Denmark, Germany, Holland,Scotland, England, Wales, Ireland, the Orkney Islands, France, Portugal, Spain, and Gibraltar; entire coast ofMediterranean Sea but not in Black Sea; Morocco, Madeira, Azores, possibly the Gulf of Guinea, and off South Africa(Western Cape). Indo-West Pacific: South-central Indian Ocean from South Africa (Eastern Cape and possiblyKwaZulu-Natal) eastward to between Prince Edward and Crozet Islands, between Kerguelen and St. Paul Islands, and thesouthern coast of Australia (southern Western Australia and South Australia, Victoria, Tasmania, New South Wales andsouthern Queensland), New Zealand (including Stewart Island). Subantarctic waters off South Georgia, Marion, Prince andKerguelen Islands. Eastern South Pacific: southern Chile south to Cape Horn.Habitat: A common littoral and epipelagic shark, most abundant on the continental offshore fishing banks but also found farfrom land in ocean basins and occasionally close inshore. It was recently caught at the mouth of a brackish estuary inArgentina but does not penetrate fresh water. This shark usually occurs in cold water, less than 18° and down to 1°C, but wasonce recorded in water 23°C. It does not occur in equatorial seas as far as is known. The porbeagle ranges in depth from thesurface and inshore waters less than 1 m deep down to at least 700 m.Biology: The porbeagle is described as very active and strong-swimming in pursuit of prey or when viewed by diversunderwater but does not engage in spectacular leaps like the shortfin mako (Isurus oxyrinchus) when hooked. Behaviourand sociobiology are poorly known, and most data available has a fisheries context.The porbeagle is found at the surface down to the bottom, singly and in schools and feeding aggregations. Porbeagles maycome inshore and to the surface in summer, but will winter offshore and beneath the surface. Fisheries catches in Europeindicate that the porbeagle has populational segregation by size (age) and sex.Porbeagles of the western North Atlantic seem to constitute a single stock that undertakes extensive migrations betweensouthern Newfoundland (Canada) in summer to at least Massachusetts (USA) in the winter. Longterm tagging data suggestthere is no mixing between this population and that of the eastern North Atlantic.Porbeagles breed on both sides of the North Atlantic, off the Atlantic coast of Europe and the British Isles, where femaleshave embryos during most of the year except July through September, and off North America from Massachusetts to Maine,where females can be found with young at all times of year. Young are apparently born in the spring off Europe, in eitherwinter-spring or late summer off North America, and probably from April to September (peak June-July [winter]) in theSouthern Hemisphere. Mating in European waters occurs in late summer, and breeding there probably occurs every year.An extended mating period seems to exist for Southern Hemisphere populations around Australia and New Zealand.

124 FAO Species Catalogue for Fishery Purposes No. 1The porbeagle is ovoviviparous and a uterine cannibal (oophagous), with litters of 1 to 5 young but the majority of litters areof four young. The foetuses grow enormously by feeding on fertilized eggs, and develop grotesquely expanded abdomensand branchial regions. Small porbeagle embryos posses fang-like functional teeth to tear open egg capsules and release thecontained ova; the fangs are shed at 34 to 38 cm FL (Francis and Stevens, 2000). The gestation period has been estimatedat about 8 or 9 months for North Atlantic and South Pacific populations. The length of the entire reproductive cycle is notknown. Pupping and nursery areas may be in continental waters, but are little-documented. In the western North Atlantic,mating is believed to take place off southern Newfoundland.Tag-recapture data and tetracycline injected sharks at liberty have been used to validate age determinations for porbeaglesin the western North Atlantic up to age 10, but longevity could be as much as 30 to 45 years (Natanson, Mello and Campana,in press). Preliminary studies in this region suggest that males mature at about 175 cm FL (~ Age 7) and females at around212 cm FL (~ Age 14) (Campana et al. 1999). Newborn porbeagles grow an estimated 15 to 20 cm per year (FL) during thefirst three years of life in the South Pacific. Prior to the intensive fishery that greatly reduced the numbers of this shark inEuropean waters, the annual mortality for the species was an estimated 18% under low human exploitation and probablyminimal predation pressure from other species. Recent research in Atlantic Canada indicates that the instantaneous naturalmortality rate of porbeagles is about 0.1.This shark is a proverbially voracious feeder on small to moderate-sized pelagic schooling fishes, including mackerel(Scomber, Scombridae) and pilchards and herring (Clupeidae), but also feeds on demersal fishes including various gadoidssuch as cod, haddock, cusk, whiting (Gadidae) and hake (Merlucciidae), icefishes (Channichthyidae), and John dories(Zeidae). Chondrichthyan prey include dogfish (Squalus acanthias, Squalidae) and tope sharks (Galeorhinus galeus,Triakidae). Cephalopod prey includes squid and cuttlefish. It will scavenge hooked fishes including cod from longlines.Predators of the porbeagle are little known (apart from humans). A small specimen from Argentina had tooth markssuggestive of a carcharhinid, perhaps Carcharhinus brachyurus, but it is uncertain if these were from a predation bout oragonistic encounter. The white shark and orca are obvious candidates for porbeagle predators, but records of predation byeither on porbeagles are not known to the author.Size: Maximum total length 300+ cm, possibly to 370 cm but most smaller; size at birth between 60 and 75 cm TL (69 to80 cm TL in South Pacific); males maturing at about 150 to 200 cm TL (196 cm TL in the western North Atlantic) and reachingat least 262 cm; females maturing at about 200 to 250 cm TL (with one reported at only 152 cm), to possibly 370 cm (withmost less than 300 cm). Females mature at about 237 cm TL in the western North Atlantic and at about 185 to 202 cm TL inthe South Pacific. There are several morphometric and L-W equations for porbeagles:Kohler, Casey and Turner (1995): W(kg) = 1.4823 x 10 -5 x FL(cm) 2.9641where: FL(cm) = 1.7939 + 0.8971 x TL(cm) (n = 13)(n = 15, western North Atlantic)Campana et al. (1999): W(kg) = 0.5 x 10 -4 x FL(cm) 2.713(n = 286, western North Atlantic)where: FL(cm) = 0.99 + 0.885 x TL(cm) (n = 361)Campana et al. (1999): FL(cm) = 4.96 x IDL(cm) 0.901 (n = 358)Campana et al. (1999): FL(cm) = 1.7 + 1.12 PCL(cm) (n = 360)Francis and Stevens (2000): PCL(cm) = –1.366 + 0.907 FL(cm)(n = 866, FL = 61 to 223 cm, New Zealand)Francis and Stevens (2000): TL(cm) = 4.165 + 1.098 FL(cm)(n = 173, FL = 63 to 180 cm, Australia)Interest to Fisheries and Human Impact: This species has been heavily fished commercially and utilized for humanconsumption in the temperate North Atlantic and the Mediterranean, but is also caught as bycatch in the SouthernHemisphere where it is the second most common shark as bycatch of the New Zealand longline fishery. World catches ofporbeagles have been reported to FAO by a number of countries, including Canada, Denmark, the Channel and FaeroeIslands, France, Germany, Iceland, Malta, New Zealand, Norway, Portugal, Spain, Saint Pierre and Miquelon Islands,Sweden, the United Kingdom, and the United States. World catches per annum ranged from 346 to 9 674 t from 1951through 1997 (mean 2 102 t) with 1 736 t reported in 1997 (FAO FishStat Plus database, 2000). The major peak was in themiddle 1960s and was followed by a declining trend with peaks and declines in the 1970s and 1980s to a level between1 000 and 2 500 t in the 1990s.Stocks in the North Atlantic have shown signs of serious overfishing in the form of greatly declining catches. Scandinavianfishers have caught porbeagles since the early nineteenth century, but only intensively during the twentieth century. Norwayand to a lesser extent Denmark have been the principle fishers of porbeagles in the North Atlantic. Norwegian catches havevaried wildly during the twentieth century, increasing from 279 t in 1926 to 3 884 t in 1933, then declining steadily to lowlevels during the second world war. Intensive fishing resumed in 1945 and peaked at 2 824 t in 1947, but then steadilydeclined to 207 t in 1970 and only 25 t in 1994. Porbeagles became scarce off Europe and the Norwegian fishery spread tothe western North Atlantic, but eventually the fishery shifted to other species such as shortfin mako and swordfish. Smallregulated catches exist at present for Norway (200 t allocated in European Community waters per year) and New Zealand,with the species protected in United States waters and regulated in the European Community. Canadian catches of

Sharks of the World, Vol. 2 125porbeagles were less than 100 t/y until 1990 but landings increased in 1992 and catches have oscillated around 1 300 t/ysince 1994. The western Atlantic stock is currently considered overexploited, with declining catch rates, and a fishingmortality beyond the replacement level. A Canadian management plan that limits the number of licenses, types of gear,fishing areas and seasons, prohibits finning, and restricts recreational fishing to catch-and-release only, has been in forcesince 1995. A TAC of 1 000 t/y was introduced in the commercial fishery for the period 1997-1999 pending better scientificinformation about resource status. In the past porbeagles were considered a nuisance to commercial fishermen becausethey wrecked light gear set for bony fishes (such as cod nets) and bit fish off hooks, but probably not so much at present withgreatly depleted porbeagle stocks and decimated stocks of some of its prey species (including cod).A considerable bycatch fishery for porbeagle by Japanese longliners and probably the pelagic fishing fleets of othercountries has existed in the southern Indian Ocean and probably elsewhere in the Southern Hemisphere. The catch ispoorly known and may be little-utilized except for fins. It has figured as complementary bycatch (fins utilized) of theJapanese longline fishery for southern bluefin tuna off Tasmania. It is used fresh and dried-salted for human consumption;for oil and fishmeal for fertilizer; and for fins for shark-fin soup. The species is primarily caught with pelagic longlines; alsopelagic and bottom trawls, handlines and gill nets. It has recently showed up as bycatch of demersal longlines forPatagonian toothfish (Dissostichus eleginoides, Nototheniidae) in the southern Indian Ocean. Statistics for the SouthernHemisphere porbeagle fishery are only reported to FAO by New Zealand (21 t in 1997), which suggests that the southerncatch is largely unreported.This species has been described as a relatively less active game fish compared to the shortfin mako and white shark.However, the porbeagle has been regularly sought by sportsfishing anglers in the United Kingdom, Ireland and the UnitedStates and is a strong fighter (especially on light tackle from a small boat). It is listed as a record game fish by theInternational Game Fish Association.The porbeagle seldom if ever bites people in the water or boats (unlike its close relatives the shortfin mako and whitesharks). An older anecdotal account mentions a provoked encounter by a porbeagle that leapt at and bit a piece of clothingfrom a fisherman who was attempting to capture it. A swimmer was reported as being bitten by a ‘mackerel shark’, but thiswas not confirmed and could have resulted from mistaking a white shark or shortfin mako for a porbeagle or mackerel shark.Recently adult porbeagles have been filmed underwater making fast rushes at divers servicing oil platforms in the NorthSea, with the sharks sometimes brushing the divers and making light contact without hurting them (I. Fergusson, pers.comm.). The motivation of this activity is uncertain but is apparently nonpredatory and possibly agonistic or exploratory. Tothe writer’s knowledge porbeagles have not figured in ecotouristic diving, nor have they been kept in captivity.The conservation status of the porbeagle is of major concern because of the drastic decline in catches from targetedfisheries in the North Atlantic and continuing exposure of the species to intensive high-seas pelagic longline fisheries (withfinning and capture trauma contributing to mortality) wherever it occurs. North Atlantic fisheries are relativelywell-documented and under regulation, but not those of the Southern Hemisphere with the exception of New Zealand.Local Names: Mackerel shark, Common porbeagle, Porbeagle shark, Atlantic mackerel shark, Common Atlantic mackerelshark, Salmon shark, Atlantic porbeagle, Swordfin, Blue dog, American porbeagle, Beaumaris shark, Blue shark,Bottle-nosed shark (England and USA); Le squale nez, Le lamie long nez, Lamie, Nez, Touille, Touilele boeuf taupe, Requinlong nez, Loutre de mer, Nas llarg, Melantoun (France); Neushaai (Holland); Sillhaj or Herring shark, Sildehaaen(Denmark); Morgi mawr (Wales); Haabranden, Haamar (Sweden); Smeriglio, Lamna smeriglio, Isuro muso acuto, Cagnia,Smeriglio, Sorglio pisci tunnu, Pisci cani, Cani di mer (Italy); Psina atlantska (Adriatic); Calderon, El marrago, Ludia,Marraco, Marraquet, Marrraix, Marrajo, Taulo (Spain); Anequim, Arrequim, Marracho, Sardo (Portugal); Marracho,Porbeagle (Azores); Requim, Nequim (Madeira); Akula sel devaia (Russia); Haringhaai (South Africa).Literature: Shann (1911, 1923); Garman (1913); Lahille (1928); Fowler (1936, 1941); Whitley (1939, 1940); Bigelow andSchroeder (1948); Smith (1949); Aasen (1961, 1963); Farquhar (1963); Garrick and Schultz (1963); Templeman (1963);Kato, Springer and Wagner (1967); Nakaya (1971); Stevens (1973); Baldridge (1974); Schwartz and Burgess (1975);Sadowsky and Amorim (1977); Svetlov (1978); Duhamel and Ozouf-Costaz (1982); Stevens, Dunning and Machida (1983);Compagno (1984, 1990b, c); Quero (1984); Duhamel and Compagno (1985); Sadowsky, Arfelli and Amorim (1985); Paustand Smith (1986); Stevens (1990); Last and Stevens (1994); Santos, Porteiro and Barreiros (1997); Lucifora and Menni(1997); O’Boyle et al. (1998); Castro, Woodley and Brudek (1999); Campana et al. (1999); Francis and Stevens (2000);Natanson, Mello and Campana, (in press); I. Fergusson, (pers. comm.).click for next page

click for previous page126 FAO Species Catalogue for Fishery Purposes No. 12.3 Order ORECTOLOBIFORMES - Carpet sharksOrder: Orectolobiformes: Compagno, 1973, J. Linn. Soc. (Zool.), 53, suppl. 1: 28; also Applegate, 1974, J. Mar. Biol. Ass.India, 14(2): 743.Number of Recognized Families: 7.Synonyms: [Part] 1 Squali, Abtheilung [Division] 1: Müller and Henle, 1838d: 3. [Part] 1 Squali, Abtheilung [Division] 2,Unterabtheilung [Subdivision] 3: Müller and Henle, 1839: 66. Ordo Plagiostomi, Subordo Squalini, Sectio Proktopterides,Tribus Dinotopterini: Bleeker, 1859: xi. Order Squali, Suborder Squali: Gill, 1862b: 394, 396. Order Squali, Suborder Galei:Gill, 1872: 22, 23. Order Plagiostomi diplospondyli, Suborder Plagiostomi asterospondyli, Group 1 Scyllia: Hasse, 1879: 52.Order Plagiostomi diplospondyli, Suborder Plagiostomi asterospondyli, Group 2 Scylliolamnidae: Hasse, 1879: 51. OrderSelachii, Suborder Asterospondyli: Woodward, 1889: 157. Order Asterospondyli, suborder unnamed: Gill, 1893: 130;Fowler, 1941: 4, 13; Smith, 1949: 37, 39. Order Asterospondyli, Suborder Galei: Jordan and Evermann, 1896: 19, 21. OrderEuselachii, Suborder Pleurotremata, Division Galeoidei: Regan, 1906a: 723. Order Selachii, Group 2, Division B,Subdivision 1, Suborder Scylliodei: Goodrich, 1909: 148. Order Pleurotremata, Suborder Galeoidei: Engelhardt, 1913: 97.Order Plagiostoma, Suborder Antacea, “Group” Catuloidei: Garman, 1913: 11, 12. Order Plagiostoma, Suborder Antacea,“Group” Isuroidei: Garman, 1913: 10, 12. Order Euselachii, Suborder Galei, [Series] Scyllioidei: Jordan, 1923: 97. OrderPlagiostomi, Suborder Galeiformes: Lozano y Rey, 1928: 280. Order Galea, Suborder Isurida, Superfamily Orectoloboidea:White, 1936: 4; White, 1937: 36, tab. 1. Order Euselachii, Suborder Scylliformes: Bertin, 1939a: 9. Order Lamniformes,Suborder Lamnoidei: Berg, 1940: 137; Berg and Svetovidov, 1955: 65; Patterson, 1967: 670; Lindberg, 1971: 8, 257;Nelson, 1976: 33; Nelson, 1984: 51. Order Euselachii, Suborder Galeoidei, Superfamily Orectoloboidea: Whitley, 1940:68-69. Order Selachii, Suborder Galeoidea: Romer, 1945: 576; Bigelow and Schroeder, 1948: 77, 95; Romer, 1966: 350.Order Lamnoidea, Suborder Galeoidea: Schultz and Stern, 1948: 224. Order Lamnida, Suborder Lamnina: Matsubara,1955: 1-789. Order Galeiformes, Suborder Isuroidei: Arambourg and Bertin, 1958: 2030. Order Pleurotrema, SuborderGaleoidea: Norman, 1966: 7. Order Carchariida, Suborder Carchariina, Superfamily Orectolobicae: Fowler, 1967a: 89.Order Carchariida, Suborder Carchariina, Superfamily Lamnicae: Fowler, 1967a: 104. Order Squatinida, SuborderGinglymostomatoidei: Glikman, 1967: 216. Order Euselachii, Suborder Galeoidei: Blot, 1969: 702-776. OrderPleurotremata, Suborder Galeiformes: Budker and Whitehead, 1971: 5, tab. 2. Order Carcharhiniformes: Rass andLindberg, 1971: 304; Gubanov, Kondyurin and Myagkov, 1986: 3, 61. Order Orectolobiformes: Chu and Meng, 1979: 114,tab. 2; Compagno, 1984: 165; Cappetta, 1987: 26, 71; Compagno, 1988: 382; Eschmeyer, 1990: 435; Nelson, 1994: 45; deCarvalho, 1996: 55; Shirai, 1996: 32; Eschmeyer, 1998: CD-ROM. Order Orectolobiformes, Suborder Orectoloboidei:Applegate, 1974: 74. Order Orectolobiformes, Suborder Parascylloidei: Applegate, 1974: 749. Order Orectolobiformes,Suborder Rhincodontoidei: Applegate, 1974: 744. Order Orectolobiformes, Suborder Rhincodontoidea: Chu and Meng,1979: 114, tab. 2. Order Orectolobiformes, Suborder Orectoloboidea: Chu and Meng, 1979: 114, tab. 2. OrderGaleomorpha, Suborder Heterodontoidea: Carroll, 1988: 598.FAO Names: En - Carpet sharks; Fr - Requins-tapis; Sp - Tiburones tapiceros and Gatas nodrizas.Field Marks: Moderate-sized to gigantic, sluggish, demersal to active sharks without nictitating eyelids, with barbels andnasoral grooves, nostrils connected with mouth, short mouths that end in front of eyes, usually without enlarged anteriorteeth and without a gap or small intermediate teeth between anteriors and lateral teeth on each side of the upper jaw, fivenarrow to broad gill openings, two spineless dorsal fins and an anal fin.Diagnostic Features: Head conical to greatly depressed, expanded laterally or not. Snout short to very short, truncated toflattened-conical, not greatly elongated nor blade-like and without lateral teeth or rostral barbels. Eyes usually dorsolateralon head but lateral in a few derived taxa (Stegostoma, Nebrius and Rhincodon), without nictitating lower eyelids orsecondary lower eyelids, but with shallow subocular pouches in some taxa; upper eyelids not fused to eyeball. Spiraclessmall to large, close behind and about opposite level of eyes. Five pairs of gill openings present on sides of head, with theposteriormost two to four above pectoral-fin origins. Nostrils longitudinal on snout, usually with barbels (rudimentary inRhincodon), circumnarial grooves primitively present around incurrent aperture but absent in some derived taxa; nasoralgrooves present and connecting excurrent apertures of nostrils with mouth; anterior nasal flaps short to elongate andreaching mouth. Mouth small to large, usually subterminal (terminal in Rhincodon), arched to nearly transverse and short,ending in front of eyes. Labial furrows well-developed on both jaws. Teeth weakly to strongly differentiated along the jaws,with (Orectolobidae) or usually without enlarged anterior teeth and without enlarged molariform posterior teeth; without agap or small intermediate teeth between anterior and lateral teeth in the upper jaw; teeth primitively with orthodonthistological structure but osteodont in Ginglymostoma and Nebrius. Trunk cylindrical to depressed and somewhat ray-like(Orectolobidae). Caudal peduncle with or without lateral dermal ridges or lateral keels on caudal peduncle. Dermal denticlescovering entire body, not enlarged as thorns or spines. Pectoral fins small to large, somewhat expanded and ray-like in someOrectolobidae, without triangular anterior lobes that cover the gill slits. Pectoral girdle (scapulocoracoid) fairly high,U-shaped, without a medial joint, and with superscapulae directed posterodorsally and not contacting vertebral column.Pectoral-fin skeleton primitively tribasal (dibasal in some taxa), with propterygium in contact with radials and metapterygiumwithout a proximal segment; pectoral fins primitively aplesodic, with radials confined to the fin bases, but plesodic in derivedtaxa and supporting the fin webs; radial count 14 to 28, with 2 to 10 segments. Pelvic fins small to large, with vent usuallycontinuous with their inner margins (may be separate in Orectolobidae). Claspers generally with siphons in the abdomen butwithout clasper sacs; clasper glans with a pseudosiphon (sometimes absent), cover rhipidion (sometimes reduced),rhipidion (sometimes absent), exorhipidion, and often lateral clasper spurs or spines; dorsal and ventral marginals of clasper

Sharks of the World, Vol. 2 127skeleton rolled into a tube for the clasper canal. Two spineless dorsal fins present, with origin of first dorsal fin varying fromanterior to pelvic-fin origins to behind pelvic-fin insertions; dorsal-fin skeleton with segmented radials but without segmentedbasal plates. Anal fin present. Caudal fin with a long dorsal lobe and the ventral lobe strong to absent; vertebral axis weaklyto strongly elevated into the dorsal caudal-fin lobe (heterocercal caudal fin). Vertebral calcification strong, secondarycalcification usually in form of strong branched radii in intermedial spaces although sometimes wedge-shaped and solid(Parascylliidae), intermedialia sometimes with annular rings (Rhincodontidae), usually without diagonal lamellae in thebasal spaces (present in Rhincodon and Parascyllium). Total vertebral count 117 to 243, precaudal vertebrae 69 to 138.Neurocranium with a short to rudimentary monopodal rostrum consisting of a ventral rostral cartilage without ventral keel,rostral space open dorsally; nasal capsules trumpet-shaped and more or less flattened, without subnasal fenestrae (basalcommunicating canals) or antorbital cartilages; orbits with incomplete preorbital walls, usually strong supraorbital crests(greatly reduced in Parascylliidae), usually strong suborbital shelves (greatly reduced in Parascylliidae), separate foraminafor superficial ophthalmic nerves but not for hyomandibular nerves, and incomplete postorbital walls without lateralcommissures for lateral head vein; occipital condyles low, occipital hemicentrum present between them. Jaws short tomoderately elongated, upper jaws (palatoquadrates) with low, ridge-like orbital processes that articulate with nasal capsulesand orbits in horizontal grooves contacting ethmoid region, basal plate and suborbital shelves; orbital processes notpenetrating supraorbital crests. Hyobranchial skeleton with moderately broad, short to elongated basihyoid; posterior twopharyngobranchials and last epibranchial fused into a yoke-shaped element. Head muscles include vertical broadpreorbitalis; short levator palatoquadrati and separate first dorsal constrictors that do not extend behind the postorbitalprocesses, adductor mandibulae muscles segmented into two or three groups of divergently biased fibres, not notchedanteriorly for mouth gape; no craniomandibular muscle between the lower jaw and orbital walls; no mandibulocutaneousmuscle between upper jaw and skin; and no postocular eyelid muscles (an anterior palpebral depressor muscle present inParascylliidae). Intestinal valve of conicospiral or (usually) ring type, with 6 to 74 turns. Reproduction oviparous in somespecies, which lay eggs in oval cases; other species are ovoviviparous (aplacental viviparous), with foetal nutriment from theyolk sack or from uterine cannibalism (oophagy or egg-eating in Nebrius), but without placental vivipary or nutritivetrophonemata.Distribution: Circumglobal in warm-temperate and tropical seas, absent or marginal in cold-temperate areas and notreaching cold boreal and subantarctic waters. Most species occur in tropical seas and are most diverse in the westernPacific. The whale shark is circumglobal, and some large carpet sharks have wide ranges in the Atlantic and eastern Pacific(Ginglymostoma), or the Indo-West Pacific (Nebrius and Stegostoma). Most carpet sharks have more limited ranges in theIndo-West Pacific, with several species confined to Australia and New Guinea and a few occurring only off Taiwan (Provinceof China) or Japan.Habitat: Carpet sharks or orectoloboids occur in a variety of marine habitats from shallow open and enclosed bays, rockyand coral reefs, estuaries, and sandy beaches on the continental shelves to the outer shelves, uppermost slopes, and theepipelagic zone (Rhincodontidae). They range in depth from the intertidal to the outer shelves and exceptionally between200 and 435 m on the upper slopes (Parascylliidae) and in the epipelagic zone at the surface (whale shark, Rhincodon).Although many species are found in shallow inshore waters only a few species of Hemiscylliidae may penetrate the lowerbrackish reaches of rivers but are not definitely recorded from freshwater rivers and lakes. No carpet sharks are specializeddeep-slope or oceanic species.Biology: Carpet sharks are small to gigantic, rather varied sharks that are mostly small, benthic, and sluggish, while largerspecies are generally more active swimmers. Morphotypes (Compagno, 1990a) include generalized benthic and littoralspecies (Ginglymostomatidae and Brachaeluridae), elongated leptobenthic bottom-dwelling species (Parascylliidae andHemiscylliidae), squatinobenthic depressed specialists (Orectolobidae), and a macroceanic, filter-feeder (Rhincodontidae).The squatinobenthic wobbegongs are ambush-hunters that take relatively large prey, swallowed whole, and parallel themostly allopatric angel sharks (Squatinidae). The prey of orectoloboids includes microscopic zooplankton, small tomoderate-sized bony fishes, smaller chondrichthyans, crabs, lobsters, shrimp, octopuses, cuttlefish, squid, gastropods,bivalves, sea urchins, sea anemones, and corals. Information on movements is limited or absent for most species; the whaleshark is highly migratory, and seasonally visits favoured areas including concentrations of food (reefs with plankton blooms.)The whale shark is apparently social and forms aggregations, but the sociobiology of most species is poorly known. Modesof reproduction include oviparity (egg-laying, in Parascylliidae, Hemiscylliidae, and Stegostomatidae), ovoviviparity oraplacental vivipary (Brachaeluridae, Orectolobidae, Rhincodontidae, Ginglymostoma), and uterine cannibalism or cannibalvivipary in the form of egg-eating or oophagy (Nebrius).Interest to Fisheries and Human Impact: Several families of orectoloboids are minor to important fisheries sharks incoastal and oceanic waters, particularly members of the Orectolobidae, Hemiscylliidae, Stegostomatidae andGinglymostomatidae. These are regular components of targeted shark fisheries and as bycatch of other fisheries targetingteleost fishes or marine invertebrates. Some species are discarded bycatch of demersal fisheries (Parascylliidae), while thewhale shark is targeted by specialized fisheries. Orectoloboids are caught in bottom trawls, in fixed and possibly pelagic gillnets, in fish traps, on bottom longlines, with harpoons, and with hook-and-line and rod-and-reel. Several species are usedfor human consumption; the flesh of some species is excellent, and large fins are of high value in the oriental soup-fin trade.Whale sharks were formerly caught by very small artisanal fisheries for local use, but values of whale shark fins, flesh andother products for the international market have soared during the last decade and have encouraged intensive targetedfisheries in certain countries (India, Philippines, and Taiwan (Province of China)). Several inshore and offshore species arecaught by sportsfishing anglers.

128 FAO Species Catalogue for Fishery Purposes No. 1Wobbegongs and larger nurse sharks (Ginglymostomatidae) rarely bite people. Whale sharks occasionally butt fishingboats, but are more often run down and injured or killed by ships.Whale sharks, nurse sharks, wobbegongs, and zebra sharks are currently sought by ecotouristic divers and film-makers inthe tropics, with whale sharks the subject of dedicated dive tours in the Indo-Pacific from South Africa to Costa Rica.Orectoloboids as a group are very hardy in captivity and are among the most important aquarium sharks, both for publicviewing and for home aquaria. Most species have been kept in captivity, with the wobbegongs (Orectolobidae), bamboosharks (Hemiscylliidae), nurse sharks (Ginglymostomatidae), and zebra sharks (Stegostomatidae) being the most popularorectoloboids for public aquaria and oceanaria. Whale sharks are kept in a few Japanese oceanaria with sufficiently hugetanks to accommodate them.The diversity of orectoloboid sharks is greatest in inshore and offshore continental waters of the tropical Indo-West Pacific,where intensive fisheries occur. The conservation problems facing the whale shark are the most urgent and widelypublicized, but most species of orectoloboids are taken in targeted and bycatch demersal fisheries and several species arecoral-reef dwellers that are adversely affected by reef destruction. Some species have limited geographic and bathymetricdistributions in continental inshore and offshore tropical waters (some Parascylliidae, Orectolobidae, Brachaeluridae andHemiscylliidae) and are at potential risk from habitat degradation and fisheries. Fisheries statistics are unavailable foralmost all species, including catches of carpet sharks for the aquarium trade. The whale shark is included on the IUCN RedList and protected off Belize, Honduras, the Philippines, and the USA (east coast) while nurse sharks (Ginglymostoma) areprotected off the east coast of the United States.Remarks: The concept of the orectoloboids as a discrete, monophyletic group is relatively recent and stems from the workof Regan (1906a), White (1936, 1937), Compagno (1973) and Applegate (1974). Most researchers in the nineteenth andtwentieth centuries followed Müller and Henle (1839) in placing the highly distinctive whale shark (Rhincodon) in a separate,monotypic family (Rhincodontidae or its synonyms). Müller and Henle (1838d) included orectoloboids other than the whaleshark in the carcharhinoid catshark family (Scyliorhinidae or its synonyms), and this was followed by several earlier authors(Müller, 1845; Gray, 1851; Bleeker, 1859; Dumeril, 1865; Günther, 1870; Jordan and Gilbert, 1883; Woodward, 1889;Goodrich, 1909; Bertin, 1939a).Gill (1862b, 1872) divided the orectoloboids into the families Rhinodontoidae (Rhinodontidae) for the whale shark,Ginglymostomatidae for the nurse sharks (Ginglymostoma, Nebrius), Crossorhinoidae for the wobbegongs (Orectolobus),and Scylliorhinoidae for other orectoloboids and the scyliorhinid catsharks. Gill (1862b) subdivided the Scylliorhinoidae intothe subfamilies Scylliorhininae for the catsharks and Parascylliinae, Hemiscylliinae, Chiloscylliinae, and Stegostomatinaefor other orectoloboids, and later (Gill, 1893) transferred the Stegostomatinae to the Ginglymostomatidae. Hasse (1879)named a “Group 2 Scylliolamnidae” as a family for Stegostoma, Ginglymostoma, and Orectolobus and a “FamilyChiloscyllium” for the hemiscylliid orectoloboid Chiloscyllium.Regan (1906a; followed by Engelhardt, 1913 and Berg, 1940) was the first author to associate the whale shark with otherorectoloboids in a common family Orectolobidae. White (1936, 1937) expanded Regan’s arrangement by placing the whaleshark (Rhincodontidae) and the carpet sharks (Orectolobidae) in a common superfamily Orectoloboidea. White wasfollowed by Whitley (1940), who further subdivided the Orectoloboidea into the families Hemiscylliidae, Orectolobidae,Ginglymostomatidae and Rhincodontidae in the superfamily Orectoloboidea. This arrangement and the orectoloboidfamilies and subfamilies of Gill (1862b, 1872, 1893) were revived and expanded by Compagno (1973) and Applegate(1974), who proposed a common order Orectolobiformes for all orectoloboids, and recognized the separate familiesParascylliidae, Brachaeluridae, Orectolobidae, Hemiscylliidae, Stegostomatidae, Ginglymostomatidae and Rhiniodontidae,while Applegate (1974) proposed an additional family Cirrhoscylliidae (here ranked as a synonym of Parascylliidae). Thiswas followed by several authors (Compagno, 1981b, 1982, 1984, 1988, 1999; Cappetta, 1987; Nelson, 1994; Eschmeyer,1990, 1998; de Carvalho, 1996; Helfman, Collette and Facey, 1997). Dingerkus (1986) and Shirai (1996) modified thisclassification by synonymizing the Ginglymostomatidae and Stegostomatidae with the family Rhincodontidae. Chu andMeng (1979) recognized the order Orectolobiformes with three families, Orectolobidae, Cirrhoscylliidae andRhincodontidae. Carroll (1988) recognized a suborder Orectoloboidea for the families Orectolobidae, Rhinocodontidae[sic], Hemiscylliidae and Parascylliidae.There are several alternative arrangements for the classification of orectoloboids. Orectoloboids were often included in acommon and undifferentiated ‘galeoid’ group along with the lamnoids and carcharhinoids (and occasionally theheterodontoids) as the families Orectolobidae and Rhincodontidae (Fowler, 1941; Romer, 1945, 1966; Bigelow andSchroeder, 1948; Schultz and Stern, 1948; Smith, 1949; Matsubara, 1955; Garrick and Schultz, 1963; Norman, 1966; Blot,1969; Pinchuk, 1972). Some authors associated the Rhincodontidae or equivalents with more derived lamnoids or all of thelamnoids in a common group (Bonaparte, 1838; Goodrich, 1909; Garman, 1913; Fowler, 1947, 1967a; Berg and Svetovidov,1955; Nelson, 1976, 1984). Arambourg and Bertin (1958) included all orectoloboids with the lamnoids in a common highergroup. Glikman (1964, 1967) included the orectoloboids with most of the squalomorph sharks (squatinoids, squaloids, andpristiophoroids). Jordan (1923) named a Series Scyllioidei for the orectoloboid families Hemiscylliidae, Orectolobidae,Ginglystomidae (sic.), and Rhincodontidae, which also included the carcharhinoid catsharks. Rass and Lindberg (1971) andGubanov, Kondyurin and Myagkov (1986) grouped the orectoloboids in a common, undifferentiated orderCarcharhiniformes with certain lamnoids (Cetorhinus, Megachasma), and the carcharhinoids.Continuing work on the morphology of orectoloboid sharks by the writer as an extension of previous research (Compagno,1979, 1984, 1988) supports the recognition of the Orectolobiformes as a monophyletic if morphologically varied group. The

Sharks of the World, Vol. 2 129arrangement of orectoloboid families recognized in Compagno (1973, 1984) are retained here, but one notes thatDingerkus’ (1986) cladistic classification united the more derived orectoloboids in a single family Rhincodontidae.Compagno’s (1988) cladistic analysis of orectoloboids was at variance with Dingerkus (1986) but supported the monophylyof the group of ‘higher’ orectoloboids placed in his Rhincodontidae (Stegostoma, Pseudoginglymostoma, Nebrius,Ginglymostoma and Rhincodon). A problem remains on the familial position of the genus Pseudoginglymostoma, which isretained provisionally in Ginglymostomatidae here. Alternative cladograms (Dingerkus, 1986, Compagno, 1988) suggestedthat this genus may be either the plesiomorphic sister group of all other ‘higher’ orectoloboids (which seems more likely), oralternatively is the plesiomorphic sister group of Stegostoma. Alternatives include placing Pseudoginglymostoma in its ownfamily, in the Stegostomatidae, or accepting Dingerkus’ arrangement of an expanded and morphologically diverseRhincodontidae. The writer favours the former arrangement, but reserves action for the completion of a detailed study of theanatomy of Pseudoginglymostoma and comparison with other higher orectoloboids.Key to Families:near-terminalmouth1a. Mouth huge and nearly terminal; externalgill slits very large, internal gill slits insidemouth cavity with filter screens; caudalpeduncle with strong lateral keels; caudallarge gill slitsfin with a strong ventral lobe, but without astrong terminal lobe and subterminalnotch (Fig. 80) . . . . . . . . . . family Rhincodontidaestrong ventral lobeFig. 80 Rhincodontidae1b. Mouth smaller and subterminal; externalgill slits small, internal gill slits without filterscreens; caudal peduncle without stronglateral keels; caudal fin with a weak ventrallobe or none, but with a strong terminallobe and subterminal notch (Fig. 81) . . . . . . . . . . 2subterminal mouthsmall gill slitsFig. 81 Stegostomatidaeweak ventral lobea) FRONTAL VIEWOF HEAD2a. Caudal fin about as long as rest of shark(Fig. 81) . . . . . . . . . . . . . family Stegostomatidae2b. Caudal fin much shorter than rest of shark . . . . . . . . 3Fig. 82 Orectolobidaeb) DORSAL VIEW3a. Head and body greatly flattened, headwith skin flaps on sides; two rows of large,fang-like teeth at symphysis of upper jawand three in lower jaw (Fig. 82) . . . . family Orectolobidae3b. Head and body cylindrical or moderatelyflattened, head without skin flaps; teethsmall, not enlarged and fang-like atsymphysis . . . . . . . . . . . . . . . . . . . . . . . . 4HEAD AND DETAIL OF NOSTRILFig. 83Ginglymostomatidae4a. No circumnarial lobe and groove aroundouter edges of nostrils (Fig. 83). family Ginglymostomatidae4b. A circumnarial lobe and groove aroundouter edges of nostrils (Fig. 84) . . . . . . . . . . . . . . 5HEAD AND DETAIL OF NOSTRILFig. 84

130 FAO Species Catalogue for Fishery Purposes No. 15a. Spiracles minute; origin of anal fin well inspiracle minutefront of second dorsal-fin origin,separated from lower caudal-fin origin byspace equal or greater than its baselength (Fig. 85). . . . . . . . . . . . family Parascylliidae5b. Spiracles large; origin of anal fin wellbehind second dorsal-fin origin,separated from lower caudal-fin origin byspace less than its base length (Fig. 86) . . . . . . . . 6barbelsgroovespiracle largeorigin of anal finFig. 85 Parascylliidaeorigin of anal finFig. 86 Hemiscylliidaegroove6a. Nasal barbels very long; longitudinalgroove on middle of chin; anal fin high andangular; distance from vent to lowercaudal-fin origin shorter than distancefrom snout to vent (Fig. 87). . . . . . family Brachaeluridaea) UNDERSIDE OF HEADb) LATERAL VIEWFig. 87 Brachaeluridaebarbels6b. Nasal barbels short; no groove on chin;anal fin low, rounded and keel-like;distance from vent to lower caudal-finorigin longer than distance from snout tovent (Figs 86 and 88) . . . . . . . . family HemiscylliidaeUNDERSIDE OF HEADFig. 88Hemiscylliidae2.3.1 Family PARASCYLLIIDAEFamily: Subfamily Parascylliinae Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 407, 408, 412 (FamilyScylliorhinoidae Gill, 1862). Also as Subfamily Parascylliinae Fowler, 1934, Proc. Acad. Nat. Sci. Philadelphia, 85: 238(Family Orectolobidae).Type Genus: Parascyllium Gill, 1862.Number of Recognized Genera: 2.Synonyms: Family Parascylliidae Applegate, 1974: 749. Type genus: Parascyllium Gill, 1862. Independently proposed asa family. Family Cirrhoscylliidae Applegate, 1974: 749; Chu and Meng, 1979: 37, 114, tab. 2 (independently proposed fromApplegate, 1974). Type genus: Cirrhoscyllium Smith and Radcliffe, 1913.FAO Names: En - Collared carpet sharks; Fr - Requins-carpettes; Sp - Alfombreras.Field Marks: Small sharks, superficially similar to members of the family Scyliorhinidae in their slender form, cat-like eyeswith subocular pockets, first dorsal-fin origin behind pelvic-fin bases, and fin proportions, but differing in having their mouthsentirely in front of eyes and in having narrow nasoral grooves, circumnarial grooves and folds around the nostrils, and medialbarbels not derived from the anterior nasal flaps. Their mouth and nostril structures, two spineless dorsal fins and an anal fin,anal-fin origin well ahead of second dorsal-fin origin, and minute spiracles distinguish them from other sharks.Diagnostic Features: Head narrow and somewhat flattened, without lateral flaps of skin. Snout broadly rounded to slightlypointed. Eyes dorsolaterally situated on head and with strong subocular ridges below them. Eyes with movable uppereyelids and shallow subocular pockets and ridges. Spiracles minute and much smaller than eyes, without raised externalrims; spiracles somewhat behind but not below eyes. Gill slits small, fifth gill slit overlapping fourth; internal gill slits withoutfilter screens. Nostrils with short, pointed barbels, circumnarial folds and circumnarial grooves around outer edges ofincurrent apertures. Nasoral grooves long and strongly developed. Mouth small, moderately arched, and subterminal on

Sharks of the World, Vol. 2 131head. Lower lip not trilobate and without lateral orolabial grooves connecting edge of lip with medial ends of lower labialfurrows; no symphysial groove on chin. Lower labial furrows ending well lateral to symphysis, not connected medially by amental groove nor mental groove and flap. Teeth not strongly differentiated in upper and lower jaws, with symphysial teethnot enlarged and fang-like. Tooth row count 23 to 54/22 to 49. Teeth with a strong medial cusp, a pair of short lateral cusplets,and strong labial root lobes. Teeth orthodont with a central pulp cavity and no plug of osteodentine. Body cylindrical orslightly depressed, without ridges on sides. Precaudal tail about as long as body or slightly longer. Caudal peduncle withoutlateral keels or precaudal pits. Pectoral fins small, broad and rounded. Pectoral fins aplesodic and with fin radials notexpanded into fin web. Pectoral propterygium small and fused with mesopterygium; pectoral-fin radial segments mostly 3 or(rarely) 2 or 4, and with longest distal segments less than 0.3 of length of longest proximal segments. Pelvic fins about aslarge as dorsal fins and slightly larger than anal fin, noticeably smaller than pectorals and with anterior margins 0.6 to 0.8 ofpectoral-fin anterior margins. Claspers with unique mesospurs, claws and dactyls (Compagno, 1988). Dorsal finsequal-sized. First dorsal-fin origin and insertion well behind the pelvic-fin bases. Anal fin somewhat smaller than seconddorsal fin, with broad base, angular apex, origin well ahead of second dorsal-fin origin, and insertion separated by a spacemuch greater than its base length from lower caudal origin. Caudal fin horizontally elongated and not crescentic, weaklyheterocercal and with its upper lobe hardly elevated above the body axis; dorsal caudal-fin margin less than one-fourth aslong as the entire shark. Caudal fin with a strong terminal lobe and subterminal notch but without a ventral lobe, preventraland postventral margins not differentiated but forming a continuous curve. Vertebral centra without radii, with solid or hollowintermedialia and diagonal calcified knobs present or absent. Total vertebral count 159 to 199, monospondylous precaudalcount 35 to 48, diplospondylous precaudal count 72 to 95, diplospondylous caudal count 51 to 65, and precaudal count 111to 138. Cranium narrow and not greatly expanded laterally. Medial rostral cartilage moderately long and not reduced to a lownubbin. Nasal capsules elevated and not greatly depressed but strongly fenestrated, internarial septum high andcompressed. Orbits with enlarged fenestrae for preorbital canals, medial walls greatly fenestrated around the optic nerveforamina. Supraorbital crests absent from cranium. Suborbital shelves strongly reduced. Cranial roof with a continuousfenestra from the anterior fontanelle to the parietal fossa. Basal plate of cranium with a pair of internal carotid foramina but nostapedial foramina. Adductor mandibulae muscle of jaws with two divisions. Preorbitalis muscles not extending ontoposterodorsal surface of cranium. Anterodorsal palpebral depressor, rostromandibular, rostronuchal and ethmonuchalmuscles present (Compagno, 1988) and unique to the family. Valvular intestine of conicospiral type with 6 to 9 turns.Development oviparous. Size small with adults 34 to 91 cm. Colour pattern of obscure or prominent dark saddles, with orwithout numerous small to large black or white spots, blotches and collar markings around branchial region.Distribution: These are little-known sharks of inshore to deepish temperate and tropical, continental waters of the westernPacific. One genus (Parascyllium) is confined to Australian waters while the second (Cirrhoscyllium) occurs in the SouthChina sea from Viet Nam north to Taiwan (Province of China) and Japan.Habitat: These sharks occur on the bottom from close inshore to at least 435 m deep offshore. They are found on muddy,sandy or rocky bottom.Biology: These small, harmless bottom sharks are rare to common in some areas but are poorly known biologically. Theyapparently can change colour somewhat to match the bottom type. All species are small, less than a metre long whenmature. At least some and possibly all of the species are oviparous, depositing eggs in elongated, flattened egg cases on thebottom. Food habits are little known, but probably these sharks include small fishes, crustaceans, and other bottominvertebrates in their diet.Interest to Fisheries and Human Impact: Several species are taken in bottom trawls as incidental and minimal bycatch,but utilization is probably minimal. They are harmless to people. Conservation status is uncertain, but should be monitoredas they have restricted and localized distributions and some of them occur in areas of heavy inshore and offshore fishingactivity, including demersal trawling. Some members of the family have been kept in captivity in public aquaria and makeattractive displays, particularly the spectacularly marked Parascyllium variolatum. Michael (1993) suggested that thesesharks are suitable for captive breeding programmes, being small and hardy in captivity.Local Names: Collared carpet sharks, Australian cat sharks.Remarks: Applegate (1974) proposed a separate family for the genus Cirrhoscyllium, but external and anatomical studies(Compagno, 1984, 1988) suggested that this genus is closely related to Parascyllium although readily distinguishable fromit, and that both genera are referable to a single family Parascylliidae. These sharks are remote from other orectoloboids andare distinguishable from them by their unique, carcharhinoid-like teeth with strong labial root lobes and low basal ledges,arched mouths, position of the anal-fin origin well anterior to the second dorsal-fin origin, cranial morphology, extraordinarilyderived cranial muscles, and spiral intestinal valves with few turns. The clasper morphology of parascylliids is unique andhighly derived, with a unique, medial, finger-like, spur-bearing lobe (dactyl) supported by the dorsal terminal cartilage as wellas a row of unique clasper hooks on the ventral terminal cartilage.Applegate (1974) placed the two parascylliid genera in a separate suborder of the Orectolobiformes, emphasizing theirdistinctiveness. Compagno (1988) suggested that the Parascylliidae is the plesiomorphic sister group of all otherorectoloboids. Dingerkus (1986), in contrast made Parascylliidae the sister group of Orectolobidae and Brachaeluridae only.Literature: Regan (1908a); Smith (1913); Garman (1913); Whitley (1940); Fowler (1941, 1967a); Stead (1963); Compagno(1973, 1984, 1988); Applegate (1974); Dingerkus (1986); Last and Stevens (1994); Goto and Nakaya (1996).

132 FAO Species Catalogue for Fishery Purposes No. 1Key to Genera:1a. A pair of barbels on throat (Fig. 89); colourpattern of dark saddles, no light or darkspots . . . . . . . . . . . . . . . . . Cirrhoscyllium1b. No barbels on throat (Fig. 90); colourpattern of saddles, dark spots, or dark andlight spots . . . . . . . . . . . . . . . . Parascylliumbarbelsno barbelsUNDERSIDE OF HEADFig. 89 CirrhoscylliumUNDERSIDE OF HEADFig. 90 ParascylliumCirrhoscyllium Smith and Radcliffe, 1913Genus: Cirrhoscyllium Smith and Radcliffe in Smith, 1913, Proc. U.S. Natl. Mus., 45(1997): 568.Type Species: Cirrhoscyllium expolitum Smith and Radcliffe, 1913, by original designation.Number of Recognized Species: 3.Synonyms: Genus Zev Whitley, 1927: 289. Unjustified replacement name for Cirrhoscyllium Smith and Radcliffe, 1913,thought by Whitley to be a junior synonym of Cirriscyllium Ogilby, 1908 = Brachaelurus Ogilby, 1907. Type species:Cirrhoscyllium expolitum Smith and Radcliffe, 1913, by replacement and by original designation.Diagnostic Features: Snout relatively long, narrow and pointed, head broad and flattened. A pair of cartilage-cored barbelspresent on ventrolateral surface of throat below rear corners of eyes, unique to this genus and found in no other sharks. Eyeshorizontally oval. Tooth count 23 to 32/22 to 27 in adults and subadult juveniles. Pectoral fins relatively thin and rather large,their anterior margins nearly equal to head length and to distance between bases of pectoral and pelvic fins. Vertebrae few,total count 159 to 175. Size small, adults 34 to 49 cm long. Colour pattern of dark saddles present, but no spots on body orcollar markings around gills.Remarks: Data for the three species of Cirrhoscyllium are from Smith and Radcliffe in Smith (1913), Kamohara (1943),Teng (1959b), Shiino (1972), Compagno (1984), Nakaya and Shirai (1984), Goto, Nakaya and Amaoka (1994), and Gotoand Nakaya (1996), who reviewed the genus. Compagno (1984) based keys and diagnoses for the three species ofCirrhoscyllium on Smith and Radcliffe (1913), Kamohara (1943), and Teng (1959b) as well as on examination of theholotype of C. expolitum, but noted: “It is not certain at present whether the characters used to distinguish the three speciesare valid, and if so, if the three species are synonyms. Pending further studies on the genus these species are provisionallyrecognized.”Goto and Nakaya (1996) examined a sample of 22 C. japonicum as well as the 11 paratypes of C. formosanum and theholotype of C. expolitum. They indicated that the characters used by Compagno (1984) were variable and not entirelydiagnostic. These characters include position of first dorsal-fin origin relative to the pelvic fins, position of first dorsal-finorigin relative to the snout tip and subterminal notch, position of anal-fin insertion relative to second dorsal fin, size of anal-finbase relative to the dorso-caudal space, and number of dark saddle markings on sides. They provided additionalmorphometric and colour characters (adopted here) to separate the three species. Their account suggests that C.japonicum is readily separable from the two other species, but that additional comparative material of C. expolitum and C.formosanum is desirable to determine if the morphometric and vertebral count characters of these species are valid. Thehumped back suggested by Compagno (1984) as a characteristic of Cirrhoscyllium was thought by Goto and Nakaya(1996) to be an artefact of preservation.

Sharks of the World, Vol. 2 133Key to Species (modified from Goto and Nakaya, 1996):1a. Saddle-markings bold, saddles on back over abdomendiscrete C-shaped; second dorsal-finbase usually shorter and rarely equal toanal-caudal space; precaudal vertebral count117 to 123 (Fig. 91). . . . . . Cirrhoscyllium japonicum1b. Saddle-markings diffuse, saddles on back overabdomen rounded and confluent with saddlesover pelvic fins; second dorsal-fin base equal toor longer than anal-caudal space (Fig. 92);precaudal vertebral count 108 to 117 . . . . . . . . 2Fig. 91 Cirrhoscyllium japonicumFig. 92 Cirrhoscyllium expolitum2a. Head length 3.0 times first dorsal-fin base(Fig. 92); total vertebral count 154, precaudalvertebral count 108 . . . . . . Cirrhoscyllium expolitum2b. Head length 2.3 to 2.6 times first dorsal-fin base(Fig. 93); total vertebral count 159 to 167,precaudal vertebral count 112 to 117 . . . . . .. . . . . . . . . . . . . . . Cirrhoscyllium formosanumFig. 93 Cirrhoscyllium formosanumCirrhoscyllium expolitum Smith and Radcliffe, 1913 Fig. 94Cirrhoscyllium expolitum Smith and Radcliffe in Smith, 1913, Proc. U.S. Natl. Mus., 45(1997): 568, figs 1-2, pl. 45.Holotype: US National Museum of Natural History, USNM-74603, 335 mm adult female, 21° 33’ N, 116° 13’ E, South ChinaSea between northern Luzon, Philippines, and China, 183 m. Status and correction in longitude from Howe and Springer(1993, Smiths. Contr. Zool., [540]: 7); also by author’s examination of the holotype.Synonyms: None.Other Combinations: None.FAO Names: En - Barbelthroat carpet shark; Fr - Requin-carpette á moustache; Sp - Alfombrera barbuda.DORSAL VIEW OF HEADUNDERSIDE OF HEADFig. 94 Cirrhoscyllium expolitumField Marks: Barbels on throat, nasoral grooves, mouth in front of eyes, six or possibly ten diffuse saddle marks on dorsalsurface, saddles above abdomen rounded and continuing above pelvic-fin bases, not C-shaped, head length three timesfirst dorsal-fin base.

134 FAO Species Catalogue for Fishery Purposes No. 1Diagnostic Features: Head length 3.0 times first dorsal-fin base. Anal-caudal space 7.7% of total length, 43% of headlength. Second dorsal-fin base equal to or longer than anal-caudal space. Precaudal vertebral count 108, total count 154.Colour: six to possibly ten indistinct pairs of saddle markings on sides of back and tail; an elongated rounded saddle oneach side of back between bases of pectoral and pelvic fins and extending over pelvic-fin bases.Distribution: Western North Pacific: South China Seaoff the coast of China between China and Luzon,Philippines, and in the Gulf of Tonkin off Viet Nam.Nominal from Okinawa (Uchida, 1982) but recorduncertain, possibly C. japonicum?Habitat: Outer continental shelf, South China Sea onbottom at 183 to 190 m depth.Biology: A little-known and presumably rare oruncommon tropical bottom shark. Probably oviparous,judging from large nidamental glands. Food habitsunknown.Size: The adult female holotype is 335 mm long while aVietnamese female (maturity stage unknown) is 306 cmlong (Kharin, 1987).Interest to Fisheries and Human Impact: Interest to fisheries unknown, probably taken as discarded bycatch of offshoretrawl fisheries in the area, conservation status unknown.Local Names: Higezame (Japan).Literature: Smith (1913); Fowler (1941); Teng (1959b); Compagno (1984); Kharin (1987); Goto and Nakaya (1996).Cirrhoscyllium formosanum Teng, 1959 Fig. 95Cirrhoscyllium formosanum Teng, 1959b, Taiwan Fish. Res. Inst., Keelung, Lab. Fish. Biol. Rep., (7): 1, pl. 1. Holotype:Taiwan Fisheries Research Institute, TFRI 3574, 367 mm female, off Kao-hsiung, Taiwan (Province of China), 110 m.Synonyms: None.Other Combinations: None.FAO Names: En - Taiwan saddled carpet shark; Fr - Requin-carpette chin; Sp - Alfombrera de Taiwan.TEETHDERMAL DENTICLEUNDERSIDE OF HEADFig. 95 Cirrhoscyllium formosanumField Marks: Barbels on throat, nasoral grooves, mouth in front of eyes, six diffuse saddle marks on dorsal surface, saddlesabove abdomen rounded and continuing above pelvic-fin bases, not C-shaped, head length 2.3 to 2.6 times first dorsal-finbase.

Sharks of the World, Vol. 2 135Diagnostic Features: Head length 2.3 to 2.6 times first dorsal-fin base. Anal-caudal space 6.8 to 7.7% of total length, 38 to45% of head length. Second dorsal-fin base equal to or longer than anal-caudal space. Precaudal vertebral count 112 to117, total count 159 to 167. Colour: six pairs of indistinct saddle markings on sides of back and tail; an elongated roundedsaddle on each side of back between bases of pectoral and pelvic fins and extending over pelvic-fin bases.Distribution: Western North Pacific: Taiwan Island(Province of China).Habitat: Outer shelf of Taiwan (Province of China) atdepth of about 110 m.Biology: A little-known small, tropical or subtropicalshark, biology essentially unknown. Twelve specimens(11 paratypes and the holotype) are in the TaiwanFisheries Research Institute, all collected offKao-hsiung, Taiwan (Province of China). That all theparatypes were collected in one day on longlinessuggests that the species may be or was commonalthough possibly localized.Size: Maximum 39 cm; eight males 35.2 to 37.7 cm; oneparatype male 35.9 cm TL from this series was adultaccording to Goto and Nakaya (1996), and presumablythe larger males were also mature. Females 35.2 to38.5 cm possibly adult if adult females attain a sizesimilar to C. expolitum.Interest to Fisheries and Human Impact: Interest to fisheries unknown, taken on bottom longlines off Taiwan (Province ofChina), presumably also as discarded trawl bycatch. Conservation status unknown.Literature: Teng (1959b); Compagno (1984); Goto and Nakaya (1996).Cirrhoscyllium japonicum Kamohara, 1943 Fig. 96Cirrhoscyllium japonicum Kamohara, 1943, Bull. Biogeogr. Soc. Japan, 13(17): 126, fig. 1. Holotype: 485 mm female fromMimase, Shikoku, Japan, possibly at Kochi University, Kochi City, Japan (Dr T. Abe, pers. comm.). According to Goto andNakaya (1996: 205), Kamohara (1961, Rep. Usa Mar. Biol. St. 8: 1-9, not seen) designated a neotype (Kochi University,Department of Biology, Faculty of Science, BSKU-3656, 291 mm immature female, from Mimase Fish Market) to replace theholotype, which was destroyed during the second world war. Goto and Nakaya suggested that Kamohara’s designation wasinvalid according to the International Code of Zoological Nomenclature (1985, Art. 75b[i] and 75d[1,2]), but redesignated thesame specimen as neotype.Synonyms: None.Other Combinations: None.FAO Names: En - Saddled carpet shark; Fr - Requin-carpette chat; Sp - Alfombrera japonesa.Fig. 96 Cirrhoscyllium japonicumUNDERSIDE OF HEAD

136 FAO Species Catalogue for Fishery Purposes No. 1Field Marks: Barbels on throat, nasoral grooves, mouth in front of eyes, nine well-defined saddle marks on sides of body,with C-shaped saddles on abdomen.Diagnostic Features: Head length 2.2 to 2.6 times first dorsal-fin base. Anal-caudal space 7.5 to 9.7% of total length.Anal-caudal space 45 to 62% of head length. Second dorsal-fin base usually shorter and rarely equal to anal-caudal space.Precaudal vertebral count 117 to 123. Total vertebral count 165 to 175. Colour: nine well-defined pairs of saddle-markingson sides of back and tail; a strongly marked C-shaped saddle on each side of back between pectoral and pelvic-fin basesand discrete from saddles over pelvic-fin bases.Distribution: Western North Pacific: Japan, offsouthwest coast from Shikoku and Kyushu southwest toYakushima Island and possibly the Riu-Kyu Islands.Habitat: Uppermost slope of southwestern Japan atdepths of 250 to 290 m.Biology: Biology poorly known. Apparently oviparous,as cased eggs were discovered in a 445 mm female. Amorphological study of the throat barbels of this shark(Goto, Nakaya and Amaoka, 1994) suggested that theywere mechanical sensors, without obvious chemical orother sense organs, and that they were unique to thegenus Cirrhoscyllium.Size: Maximum 48.5 cm; females immature at 25.7 to42.4 cm, adolescent at 43.1 cm, and adult at 44.5 cm,maximum 48.5 cm; males immature at 22.8 cm,adolescent at 36.6 cm, and adult at 36.7 to 40.7 cm.Interest to Fisheries and Human Impact: Interest to fisheries unknown. Possibly taken as a bycatch of trawl fisheries.Conservation status unknown.Local Names: Kurakake-zame, or Kurakakezame, Higezame, Saddled catshark (Japan).Remarks: A female specimen nominally referred to C. expolitum (Uchida, 1982) was collected off the Riu-Kyu Islands andkept in an aquarium in Okinawa where it laid eggs; it may refer to this species.Literature: Kamohara (1943); Teng (1959b); Uchida (1982); Compagno (1984); Goto, Nakaya and Amaoka (1994); Gotoand Nakaya (1996).Parascyllium Gill, 1862Genus: Parascyllium Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 408, 412.Type Species: Hemiscyllium variolatum Dumeril, 1853, by original designation.Number of Recognized Species: 4.Synonyms: Subgenus Neoparascyllium Whitley, 1939: 227 (Genus Parascyllium Gill, 1862). Type species: Parascylliummultimaculatum Scott, 1935, by original designation.Diagnostic Features: Snout relatively short, thick, and broadly rounded, head narrow and cylindrical. No barbels on throat.Eyes more elongated and slit-like. Tooth count 37 to 54/33 to 49 in adults. Pectoral fins thick, muscular, and rather small,their anterior margins much less than head length and than distance between pectoral and pelvic-fin bases. Vertebraenumerous, total count 188 to 199. Size of adults 60 to 91 cm long. Colour pattern of light or dark spots present, sometimeswith dark saddles, black blotches, and collar markings around gills.Remarks: The present arrangement of the species of Parascyllium follows Whitley (1940), Compagno (1984), and Lastand Stevens (1994) in most details.

Sharks of the World, Vol. 2 137Key to Species (after Compagno, 1984, and Last and Stevens, 1994):1a. Gill region with a prominent dark collarmarking dotted with dense white spots;body with brown blotches and small whitewhite spotsspots; fins with very prominent large blackblotches (Fig. 97) . . . . . . Parascyllium variolatum1b. Gill region with or without an indistinct toprominent dusky collar marking, with orwithout a few brown spots but lackingwhite spots; body with small to moderatelylarge brown spots but no blotches; finswithout black blotches (Fig. 98) . . . . . . . . . . . . . 2black blotchesFig. 97 Parascyllium variolatum2a. Gill region with a prominent dusky collar,sharply delimited from front of head; darkspots absent from pectoral fins (Fig. 98). . . . . . . . . . . . . . . . . . . . Parascyllium collare2b. Gill region with collar marking absent, paleor obscure, not sharply delimited fromfront of head; dark spots usually presenton pectoral fins . . . . . . . . . . . . . . . . . . . . . 3dusky collar without white spotsFig. 98 Parascyllium collare> 6 spotsFig. 99 Parascyllium ferrugineum3a. More than six spots on flanks betweendorsal fins (Fig. 99); a littoral species onthe continental shelves . . . . . Parascyllium ferrugineum3b. Less than six dark spots on flanksbetween dorsal fins (Fig. 100); adeepwater species on the continentalslopes . . . . . . . . . . . . . . . . . . Parascyllium sp. A< 6 spotsFig. 100 Parascyllium sp. AParascyllium collare Ramsay and Ogilby, 1888 Fig. 101Parascyllium collare Ramsay and Ogilby, 1888, Proc. Linn. Soc. New South Wales, ser. 2, 3: 1310. Holotype: AustralianMuseum, Sydney, AMS-I.1874, > 76 cm (30 in) TL adult male, in deep water off Port Jackson, New South Wales, Australia.Synonyms: None.Other Combinations: None.FAO Names: En - Collared carpet shark; Fr - Requin-carpette à collarette; Sp - Alfombrera collareja.Fig. 101 Parascyllium collareField Marks: Prominent dark, unspotted, sharp-edged collar mark around the gills, dusky saddleson back and tail, and sparse, large dark spots on body, tail and fins; nasal barbels, nasoral andcircumnarial grooves present; mouth in front of eyes, two equal-sized, spineless dorsal fins and ananal fin, the first dorsal-fin origin behind the pelvic-fin bases, the anal-fin origin well in front of thesecond dorsal-fin origin.UNDERSIDE OF HEAD

138 FAO Species Catalogue for Fishery Purposes No. 1Diagnostic Features: Colour: light yellowish to reddish brown with five dusky saddles on trunk and tail; a prominent dark,unspotted collar marking around gill region; no white spots on body; no bold black blotches and spots on fins; sparse,scattered, large dark brown spots on sides and fins, except for pectoral fins, no more than two or three irregular longitudinalrows of spots on sides and less than six spots on sides of tail between the dorsal fins.Distribution: Western South Pacific: confined to the eastcoast of Australia (Victoria, New South Wales, andsouthern Queensland from Gabo Island to Mooloolaba).Habitat: A little-known temperate bottom shark of theeastern Australian continental shelf, found at depths of20 to 160 m on hard bottom including inshore rocky reefsand hard-bottomed trawling grounds.Biology: Development oviparous, but detailslittle-known. Eggcase described as flattened andelongate, but details not given. Apparently common orformerly common on hard bottom at depths of 55 to128 m on trawling grounds off New South Wales.Size: Maximum about 87 cm; adult males 80 to 85 cmlong; adult females 85 to 87 cm.Interest to Fisheries and Human Impact: Interest to fisheries essentially none, not utilized but taken as bycatch bycommercial vessels, including trawlers and possibly line boats. Conservation status unknown.Local Names: Collared catshark or Cat shark, Collared carpet shark.Remarks: According to Last and Stevens (1994) this species was confused with other species of Parascyllium and issomewhat less wide-ranging than previously thought. It was recorded from Tasmania but these records are apparentlybased on P. ferrugineum.Literature: McCulloch (1911); Whitley (1940); Fowler (1941); Stead (1963); Compagno (1984); Last and Stevens (1994).Parascyllium ferrugineum McCulloch, 1911 Fig. 102Parascyllium ferrugineum McCulloch, 1911, Zool. Resul. Fish. Exper. F.I.S. “ENDEAVOUR”, 1: 7, pl. 2, fig. 2, text fig. 2.Holotype: Australian Museum, Sydney, possibly AMS E.4604 (Eschmeyer, 1998, Cat. Fish.: CD-ROM), 730 mm female,outside Port Phillip Heads, Victoria.Synonyms: Parascyllium multimaculatum Scott, 1935: 63, fig. 1. Holotype: Queen Victoria Museum and Art Gallery,Launceston, Tasmania, QVM old no. H.T.983 (mounted, missing in 1974 according to Eschmeyer, 1998: CD-ROM), 710 mmadult male, Tamar Heads, Devon Dorset, Tasmania, Australia. Synonymy after Last and Stevens (1994: 120).Other Combinations: None.FAO Names: En - Rusty carpet shark; Fr - Requin-carpette roux; Sp - Alfombrera mohosa.Fig. 102 Parascyllium ferrugineumUNDERSIDE OF HEADDORSAL VIEW OF HEAD

Sharks of the World, Vol. 2 139Field Marks: Inconspicuous dusky collar around the gills, with or without three or four dark spots; six or seven dusky saddleson back and tail; moderately sparse to dense, large to small dark brown or blackish spots on body, tail and fins, more than sixdark spots on the sides of the tail between the dorsal fins; nasal barbels, nasoral and circumnarial grooves present, mouth infront of eyes; two equal-sized, spineless dorsal fins and an anal fin, the first dorsal-fin origin behind the pelvic-fin bases, theanal-fin origin well in front of the second dorsal-fin origin.Diagnostic Features: Colour: grey-brown with six dusky saddles on trunk, tail and caudal fin; an indistinct dark, collarmarking around gill region, unspotted or with up to three or four dark spots; no white spots on body; no bold large blackblotches on fins though unpaired fins usually have small to large rounded black spots on them; moderately sparse to dense,scattered, large dark brown spots on body and fins, three or four (mainland) to five or six (Tasmania) irregular longitudinalrows of dark spots on sides, and more than six spots on the sides of the tail between the dorsal fins.Distribution: Western South Pacific: Confined to thesouth and east coasts of Australia (Western Australia,South Australia, Victoria, from Albany to Gabo Island,and Tasmania).Habitat: A little-known temperate-water shark of theAustralian continental shelf, on or near the bottom atdepths of 5 to 150 m. Off Tasmania close inshore nearrocks and river mouths. Occurs in beds of algae on reefsor in seagrass. Hides in rocky caves and ledges duringthe day.Biology: A poorly known nocturnal shark. Oviparous,lays eggs in yellow cases with long tendrils during thesummer. Feeds on bottom-dwelling crustaceans andmolluscs.Size: Maximum about 80 cm; possible hatchling about17 cm; males mature by 60 cm, adult males recorded at71 to 75 cm; adolescent female 74 cm.Interest to Fisheries and Human Impact: Interest to fisheries essentially none, not utilized but taken as bycatch bycommercial vessels, captured in bottom trawls and possibly by line gear. Conservation status unknown.Local Names: Rusty carpet shark, Rusty catshark or Cat shark, Tasmanian spotted catshark or Cat shark, Tasmaniancarpet shark, Requin-carpette tacheté, Alfombrera tasmánica.Remarks: Scott (1935) named Parascyllium multimaculatum as a distinct species for an adult female specimen ofParascyllium from Tasmania with densely scattered small dark spots. Whitley (1939, 1940) recognized this species andproposed a new subgenus, Neoparascyllium Whitley, 1939, for it because it differed from other species in having the firstdorsal “insertion” (= origin) behind the middle of the total length and by its markings (more numerous small dark brownspots). Compagno (1984) examined specimens of Parascyllium multimaculatum from Tasmania and recognized thespecies, but noted that the species differed little from P. ferrugineum and P. collare except in colour pattern and did notwarrant a separate subgenus. The morphometric character was thought to be invalid by Compagno (1984), as twoTasmanian specimens from Green’s Beach near the Tamar River mouth with numerous spots (one of these was illustrated inCompagno, 1984) varied in this character (PD1 60% TL in an adult female vs. 48% TL in the illustrated adult male). Thissuggests that the position of the dorsal fin relative to the middle of the total length may be individually variable and may alsovary according to size and even sex in this shark and in other species of Parascyllium. Compagno (1984) distinguished P.ferrugineum and P. multimaculatum by colour pattern only as follows:P. ferrugineum: Sparse, scattered, large dark brown spots on sides and fins, no more than three or four irregularrows of spots on sides.P. multimaculatum: Numerous small to large dark brown spots on sides and fins, five or six irregular rows of spotson sides.Last and Stevens (1994) synonymized P. multimaculatum with P. ferrugineum, and noted that “Tasmanian specimens [ofP. ferrugineum] have a variable but greater average density of spots which has led to their recognition as a separatespecies.” The present account follows Last and Stevens’ synonymy while noting that it is desirable to learn more about thenature of variation (including ontogenetic changes) in the colour pattern of these poorly-known sharks. Genetic comparisonsbetween Tasmanian and mainland animals are also desirable.Literature: McCulloch (1911); Whitley (1939, 1940); Fowler (1941); Scott (1961); Stead (1963); Compagno (1984); Michael(1993); Last and Stevens (1994).

140 FAO Species Catalogue for Fishery Purposes No. 1Parascyllium variolatum (Dumeril, 1853) Fig. 103Hemiscyllium variolatum Dumeril, 1853, Rev. Mag. Zool. (2) 5: 121, fig. 1. Holotype: Museum National d’Histoire Naturelle,Paris, MNHN-1004, 36 cm TL female, from “côtes de l’Australie” (= Tasmania).Synonyms: Parascyllium nuchalis McCoy, 1874: 15, pl. 2. Holotype: National Museum of Victoria, Melbourne, Australia,NMV no number (apparently lost according to Eschmeyer, 1998: CD-ROM), 84 cm adult female, Hobson’s Bay, Victoria,Australia.Other Combinations: None.FAO Names: En - Necklace carpet shark; Fr - Requin-carpette à collier; Sp - Alfombrera colarina.Fig. 103 Parascyllium variolatumField Marks: The bold, beautiful colour pattern of this shark is unmistakable: a unique, broad, dark, white-spotted collararound the gills, striking black spots or blotches on all fins, dark blotches and dense white spots on body; also, barbels,nasoral and circumnarial grooves present, mouth in front of eyes, two equal-sized, spineless dorsal fins and an anal fin, thefirst dorsal-fin origin behind the pelvic-fin bases, and the anal-fin origin well in front of the second dorsal-fin origin.Diagnostic Features: Colour: dark greyish or brown (to chocolate brown) above and below, with six indistinct dark saddlesvariably present or obscure on trunk and tail; a blackish brown, extremely conspicuous collar marking around the gill region,densely spotted with white like a necklace of pearls; body clouded with irregular dark brown blotches and closely sprinkledwith small to large white spots; bold black spots or blotches present on all fins, including prominent blotches on theprecaudal fins and black spots alternating with white on the dorsal and ventral web of the caudal fin; discrete dark spots notpresent on sides of trunk and tail.Distribution: Western South Pacific: South coast ofAustralia (Western Australia off Dongara to SouthAustralia, and Victoria off Lakes Entrance, also northcoast of Tasmania, but possibly including more than onespecies, the typical eastern form with type localityTasmania and an additional western form from WesternAustralia).Habitat: A temperate-water bottom shark of theAustralian continental shelf at depths from inshore downto about 180 m. Apparently found in a variety of habitats,including sandy bottom, on rocky reefs, in beds of kelp,and in seagrass beds, but details of its ecology arevirtually unknown.Biology: Biology little-known, probably oviparous.Common or unabundant, but nocturnal and seldom seen during the day. Juveniles hide under rocks and bottom debris inshallow water. Coloration may vary considerably, and individuals taken on different bottoms vary in coloration including lightspots and dark mottling and saddles, but little is known of the basis of these differences.Size: Maximum about 91 cm.Interest to Fisheries and Human Impact: Probably not utilized at present in fisheries. Role in aquarium trade unknown, butan obvious candidate for public display and for private aquaria. Conservation status uncertain.Local Names: Varied carpet shark, Southern catshark, Southern collared cat shark, Necklace carpet shark, Varied catsharkor cat shark, Hémiscylle tacheté.Remarks: Last and Stevens (1994) noted that “Another white-spotted form occurring off southern Western Australia may bean additional undescribed species.”Literature: McCulloch (1911); Whitley (1940); Fowler (1941); Stead (1963); Compagno (1984); Michael (1993); Last andStevens (1994).

Sharks of the World, Vol. 2 141Parascyllium sp. A Last and Stevens, 1994 Fig. 104Parascyllium sp. A Last and Stevens, 1994, Sharks Rays Australia: 118, pl. 16.Synonyms: Not named.Other Combinations: None.FAO Names: En - Ginger carpet shark; Fr - Requin-carpette beige; Sp - Alfombrera jengibre.Fig. 104 Parascyllium sp. AField Marks: Inconspicuous dusky half collar around the gills without spots, five indistinct dusky saddles on back and tail,and sparse, large dark diffuse spots and blotches on the body and fins with fewer than six on the sides of the tail between thedorsal fins; nasal barbels, nasoral and circumnarial grooves present, mouth in front of eyes; two equal-sized, spinelessdorsal fins and an anal fin, the first dorsal-fin origin behind the pelvic-fin bases and rear tips, the anal-fin origin well in front ofthe second dorsal-fin origin.Diagnostic Features: Colour: pale brownish or greyish above, lighter below, with five inconspicuous saddles on trunk andtail; inconspicuous dusky half collar without spots around the gills; no white spots on body; no bold black spots or blotches onfins; sparse, large dark diffuse spots and blotches on the body and fins, sparse dark spots of sides not in rows, with fewerthan six spots on the sides of the tail between the dorsal fins.Distribution: Southeastern Indian Ocean: Continentalslope of Australia (Western Australia between Lancelinand Bunbury).Habitat: Upper continental slope at 245 to 435 m.Biology: Virtually unknown.Size: To at least 79 cm total length.Interest to Fisheries and Human Impact: Unknown.Local Names: Ginger carpet shark.Literature: Last and Stevens (1994).

142 FAO Species Catalogue for Fishery Purposes No. 12.3.2 Family BRACHAELURIDAEFamily: Family Brachaeluridae Compagno, 1973, J. Linn. Soc. (Zool.), 53, suppl. 1: 28. Name only, with the generaBrachaelurus and Heteroscyllium assigned to the family; Applegate, 1974, J. Mar. Biol. Assoc. India, 14(2): 745.Type Genus: Brachaelurus Ogilby, 1907.Number of Recognized Genera: 2.Synonyms: Family Brachyaeluridae Eschmeyer and Bailey, 1990: 63; Eschmeyer, 1990: 435. Consistent emendation orerror for Brachaeluridae Compagno, 1973 or Applegate, 1974.FAO Names: En - Blind sharks; Fr - Requins aveugles; Sp - Tiburones ciegos.Field Marks: Small sharks with nasoral grooves, perinasal grooves, long barbels, small transverse mouths in front of eyes,symphysial grooves, dorsolateral eyes, large spiracles below and behind eyes, no lateral skin flaps on head, two spinelessdorsal fins and an anal fin, the second dorsal-fin origin well ahead of the anal-fin origin, and a short precaudal tail muchshorter than the head and body.Diagnostic Features: Head broad and somewhat flattened, without lateral flaps of skin. Snout broadly rounded. Eyesdorsolaterally situated on head and with strong subocular ridges below them. Eyes without movable upper eyelids but withsubocular pockets and ridges below them. Spiracles very large and subequal or larger than eyes, with prominent raisedexternal rims; spiracles somewhat below and behind eyes. Gill slits small, fifth gill slit close to fourth but not overlapping it;internal gill slits without filter screens. Nostrils with very long pointed barbels; circumnarial folds and circumnarial groovespresent around outer edges of incurrent apertures. Nasoral grooves long and strongly developed. Mouth small, slightlyarched and nearly transverse, subterminal on head. Lower lip not trilobate and without lateral orolabial grooves connectingedge of lip with medial ends of lower labial furrows, but with a longitudinal symphysial groove on chin. Lower labial furrowsextending medially nearly to symphysis, but not connected medially by a mental groove or mental flap. Teeth not stronglydifferentiated in upper and lower jaws, with symphysial teeth not enlarged nor fang-like. Tooth row count 32 to 34/21 to 29.Teeth with a strong medial cusp, a pair of short lateral cusplets, and weak labial root lobes. Teeth orthodont with a centralpulp cavity in crowns and no plug of osteodentine. Body cylindrical or moderately depressed, without ridges on sides.Precaudal tail shorter than body. Caudal peduncle without lateral keels or precaudal pits. Pectoral fins moderately large,broad and rounded. Pectoral fins aplesodic and with fin radials not expanded into fin web. Pectoral fin tribasal, propterygiumvery large and separate from mesopterygium and metapterygium; pectoral-fin radial segments three at most, and withlongest distal segments less than 0.3 times the length of longest proximal segments. Pelvic fins about as large as dorsal finsand much larger than anal fin, nearly as large as pectoral fins and with anterior margins 0.5 to 0.7 times the pectoral-finanterior margins. Claspers poorly known but probably without mesospurs, claws or dactyls. Dorsal fins equal-sized or withsecond dorsal fin slightly smaller than first. First dorsal-fin origin over pelvic-fin bases, insertion well behind the pelvic-finrear tips. Anal fin less than half as large as second dorsal fin, with broad base, angular apex, origin about opposite seconddorsal-fin midbase or insertion, and insertion separated by a space or narrow notch much less than base length from lowercaudal-fin origin. Caudal fin horizontally elongated and not crescentic, weakly heterocercal with its upper lobe at a low angleabove the body axis; dorsal caudal-fin margin about a fourth as long as the entire shark. Caudal fin with a strong terminallobe and subterminal notch but without a ventral lobe, preventral and postventral margins not differentiated and forming acontinuous curve. Vertebral centra with well-developed radii. Total vertebral count 117 to 142, monospondylous precaudalcount 30 to 40, diplospondylous precaudal count 35 to 50, diplospondylous caudal count 46 to 62, and precaudal count 69 to90. Cranium narrow and not greatly expanded laterally. Medial rostral cartilage moderately long and not reduced to a lownubbin. Nasal capsules elevated and not greatly depressed or fenestrated, internarial septum high and compressed. Orbitswith small foramina for preorbital canals, medial walls not fenestrated around the optic nerve foramina. Supraorbital crestspresent on cranium but not laterally expanded and pedicellate. Suborbital shelves moderately broad and not greatlyreduced. Cranial roof with isolated small frontal and parietal fenestrae but without a continuous fenestra from the anteriorfontanelle to the parietal fossa. Basal plate of cranium with a pair of stapediocarotid foramina. Adductor mandibulae muscleof jaws with two divisions. Preorbitalis muscles not extending onto posterodorsal surface of cranium. No anterodorsalpalpebral depressor, rostromandibular, rostronuchal or ethmonuchal muscles. Valvular intestine of spiral-ring type with 11 or12 turns. Development ovoviviparous. Colour pattern of a few broad darker saddles present on back in young but obscure orabsent in adults; small white spots present or absent. Size small, with adults between 52 and 122 cm but exceptionally over1 m total length; young are born at 15 to 18 cm.Distribution: Blind sharks are known from the western South Pacific, in the coastal waters of Australia from the east coastoff New South Wales and Queensland; additional records from Western Australia and Northern Territory need confirmation.Habitat: These are inshore bottom sharks of temperate and tropical continental coastal waters, ranging in depth from theintertidal down to 137 m. They commonly occur on rocky reefs, in seaweed, or on coral close inshore, sometimes in wateronly sufficient to cover them.Biology: These are small sharks that attain a total length of less than 1.3 m, with most individuals less than 80 cm long.Live-bearing (ovoviviparous or aplacental viviparous), with foetal nutrition primarily from the large, yolky eggs; foetuses havelarge yolk-sacks which are reabsorbed just before birth. Litter size 6 to 8. Known prey of these sharks include small fishes,crustaceans, squid, and sea anemones. At least one of the species is very hardy and can live a long time out of water as well

Sharks of the World, Vol. 2 143as surviving readily in captivity. The name ‘blind shark’ stems not from lack of vision but because these sharks close theireyelids when removed from the water.Local Names: Blind sharks.Remarks: There are two living and monotypic genera in this family: Brachaelurus Ogilby, 1907 and Heteroscyllium Regan,1908, which are confined to Australian inshore coastal waters. The members of this family were included in the familyOrectolobidae or Crossorhinidae until Compagno (1973) and Applegate (1974) placed them in their own familyBrachaeluridae, which has been followed by various authors including Compagno (1981b, 1982, 1984, 1988), Cappetta(1986), Eschmeyer (1990, 1998), and Last and Stevens (1994), but not Nelson (1976, 1984), Gubanov, Kondyurin andMyagkov (1986), and Carroll (1986), who retain them in the Orectolobidae.The systematic status of the genera of Brachaeluridae was uncertain until recently and was confounded by nomenclaturalproblems and doubts on the validity of the species Heteroscyllium colcloughi. Ogilby (1907) proposed his new genusBrachaelurus for Chiloscyllium modestum Günther, 1871 (equals Squalus waddi Bloch and Schneider, 1801).Brachaelurus has been uniformly recognized by subsequent authors. Ogilby (1908) proposed a second new genus,Cirriscyllium, for C. modestum while transferring Brachaelurus to his new species B. colcloughi. Regan (1908c) notedthat as the two species are apparently generically distinct, Cirriscyllium was a junior synonym of Brachaelurus Ogilby,1907 and that a new generic name, Heteroscyllium Regan, 1908, was necessary to replace Brachaelurus sensu Ogilby(1908) for the species B. colcloughi.Ogilby and McCulloch (1908), in a review of Australian Orectolobidae, adapted Regan’s arrangement of Heteroscyllium asa genus for B. colcloughi and Brachaelurus as a genus for Chiloscyllium modestum (= Brachaelurus waddi). Engelhardt(1913), Ogilby (1916), McCulloch and Whitley (1925), Whitley (1934, 1940), Bigelow and Schroeder (1948), Fowler (1967a),Compagno (1973, 1981b, 1982, 1984), and Applegate (1974) all followed this arrangement. Exceptionally and inexplicablyFowler (1967a) listed Hemiscyllium trispeculare Richardson, 1843 as a species of Brachaelurus along with B. waddi.Garman (1913) ranked Heteroscyllium as a subgenus for B. colcloughi within Brachaelurus, which also included B.modestus. Fowler (1929), White (1937), and more recently Last and Stevens (1994) placed H. colcloughi and B. modestumor B. waddi in Brachaelurus but did not recognize Heteroscyllium as a genus or subgenus. In contrast Fowler (1941)followed Ogilby (1908) in recognizing Brachaelurus as a genus based on B. colcloughi, but reduced Cirriscyllium to asubgenus of Brachaelurus for inclusion of B. waddi.Whitley (1940) provided the first published illustrations of Heteroscyllium colcloughi (lateral view and underside of head)and a short diagnosis and description of the species. According to Whitley (1940), the type and only known specimen of H.colcloughi was a young male 45.7 cm (1.5 ft) long that was no longer preserved (as noted by Ogilby, 1916) and theillustrations (Whitley, 1940, fig. 77, 78) were from sketches of “the type” by A.R. McCulloch made “some years ago”. Whitleyapparently overlooked the existence of the second, Queensland Museum syntype of H. colcloughi (QM I-965, designatedas lectotype below) as mentioned by Ogilby (1908, 1916) and Ogilby and McCulloch (1908), and which still exists. It is alsounclear from Whitley’s account if the sketches of the “type” were made from the specimen that was lost (AFAQ no. 410, seespecies account of H. colcloughi below) or from QM I-965.Unfortunately Whitley’s illustrations of the “type” of H. colcloughi are deceptive, leading Compagno (1984) to suggest thatHeteroscyllium differed from Brachaelurus by lacking a symphysial groove and by having a much longer caudal fin,smaller spiracles well behind eyes, subterminal nostrils, a more anterior anal-fin origin, and a longer anal-caudal space.Dingerkus (1986) followed Whitley (1934, 1940) in suggesting that only a single specimen of H. colcloughi existed andstated: “Except for its lack of a chin cleft, descriptions of it fall within the variability of Brachaelurus waddi. Unless otherspecimens are collected to confirm its existence, I consider Heteroscyllium colcloughi to be based on an aberrantspecimen of Brachaelurus waddi and thus synonymize Heteroscyllium colcloughi (Ogilby) under Brachaelurus waddi(Bloch et Schneider)”.Apart from the Queensland Museum syntype, an additional smaller specimen of Heteroscyllium colcloughi is preserved inthe collection of the Australian Museum, Sydney (P. Last, pers. comm.). Last and Stevens (1994) noted that both H.colcloughi and B. waddi had symphysial grooves and were valid species. They considered Heteroscyllium a synonym ofBrachaelurus on the suggestion that Compagno (1984) had primarily based the separation of the two genera on thesymphysial groove character (which is not correct). They also gave a descriptive account of H. colcloughi and an accurateillustration presumably based on the Australian Museum specimen.The Queensland Museum has over the past few decades acquired several additional specimens of Heteroscylliumcolcloughi from Moreton Bay near Brisbane, and currently has 13 catalogued lots with all stages from late foetuses to adultfemales. Several of these specimens were examined by the writer (including the surviving syntype), and compared withspecimens of Brachaelurus waddi during a visit to the Queensland Museum in July 1996. This suggests that a symphysialgroove may have been omitted from Whitley’s (1940) head sketch of H. colcloughi, which was otherwise accurate incomparison with specimens of H. colcloughi (all of which have the symphysial groove present). Also, Whitley’s lateral viewsketch shows the caudal fin too long and the spiracles are too small and too far from the eyes in comparison with specimens.With spurious characters eliminated, H. colcloughi can be separated from B. waddi in its own genus as clearlydemonstrated by Ogilby (1908) and as redefined below.

144 FAO Species Catalogue for Fishery Purposes No. 1Literature: Ogilby (1907, 1908); Regan (1908a,c); Ogilby and McCulloch (1908); Garman (1913); Whitley (1934, 1940);Fowler (1941); Bigelow and Schroeder (1948); Compagno (1973, 1984); Applegate (1974); Dingerkus (1986); Michael(1993); Last and Stevens (1994).barbels without flapsKey to Genera:1a. Barbels without expandedposterior flaps; seconddorsal fin about as large asfirst; anal-fin insertion justanterior to lower caudal-finorigin; usually brown abovewith white spots, no whitepatches on anterior marginsand webs of dorsal fins(Fig. 105) . . . . . . . . . Brachaelurusbarbels with flaps1b. Barbels with expandedposterior flaps at their midlengths;second dorsal finsmaller than first; anal-fininsertion separated fromlower caudal origin by aspace about equal to anal-fininner margin; usually greya) UNDERSIDE OFabove without white spots,HEADwhite patches on anteriormargins and webs of dorsalfins (Fig. 106) . . . . . . Heteroscylliuma) UNDERSIDE OF HEADwhite spotsb) LATERAL VIEWFig. 105 Brachaelurus waddib) LATERAL VIEWFig. 106 Heteroscyllium colcloughismall spacelarge spaceBrachaelurus Ogilby, 1907Genus: Brachaelurus Ogilby, 1907, Proc. R. Soc. Queensland, 1906, 20: 27.Type Species: Hemiscyllium modestum = Chiloscyllium modestum Günther, 1871, by original designation.Number of Recognized Species: 1.Synonyms: Genus Cheloscyllium Ramsay, 1880: 97, in part. Name only, apparent error for Chiloscyllium Müller andHenle, 1837. Genus Cirriscyllium Ogilby, 1908: 2, 4. Type species: Chiloscyllium modestum Günther, 1871, by originaldesignation. Unjustified replacement for genus Brachaelurus Ogilby, 1907.Field Marks: See species account of Brachaelurus waddi.Diagnostic Features: Head short and flat in adults, head about 19% of total length, maximum head height about 0.6 timeshead width; head broadly arched in dorsoventral view; snout very short, prenarial snout about 1.5% and preoral snout about3.2% of total length; snout bluntly rounded in lateral view, with ventral surface of prenarial snout nearly vertical. Eye smalland ovate with length about 1.5% of total length; eyes elevated above level of head. Spiracles horizontally situated andovate, about opposite rear ends of eyes. Nostrils nearly terminal on snout; nasal barbel without an expanded posterior flap atits midlength. Anal-caudal space virtually obsolete and much less than anal-fin inner margin. Denticles large and rough. Firstdorsal fin with origin usually slightly posterior to middle of pelvic-fin bases; apex posterior to insertion; free rear tip bluntlyangular or rounded. Dorsal fins about equal-sized, with similar height and base length. Second dorsal-fin apex posterior toinsertion; rear tip bluntly angular. Anal-fin origin about under second dorsal-fin insertion or under last fourth of seconddorsal-fin base; anal-fin free rear tip extends well behind dorsal caudal-fin origin. Total vertebral count 140 to 142, precaudalcount 88 to 90, monospondylous precaudal count 39 to 40, diplospondylous precaudal count 49 to 50. Colour: backgroundcolour of the dorsal surface dark brown, with scattered white spots on fins and body; adults with faint saddles but withoutwhite blotches on anterior webs of dorsal fins; young without black blotches on posterior dorsal-fin webs and along caudalbase but with dark saddles with very narrow transverse light lines between them.

Sharks of the World, Vol. 2 145Brachaelurus waddi (Bloch and Schneider, 1801) Fig. 107Squalus waddi Bloch and Schneider, 1801, Syst. Ichthyol.: 130. No type material. Australia?? Whitley, 1934, Mem.Queensland Mus. 10(4): 182 considered S. waddi to be the earliest name for this species, but the description could applyalso to Chiloscyllium punctatum Müller and Henle, 1838. Bloch and Schneider mention an illustration of S. waddi made byDr Latham, but unfortunately did not reproduce it in their plates. Whitley (1934) stated: “The type painting was evidentlyprepared from a specimen collected near Sydney, New South Wales, by Dr Latham, and the description most nearly appliesto the ‘blind shark’, which was later called Chiloscyllium modestum by Günther and C. furvum by Ramsay.” However,Whitley, 1941, Fish. Australia 1: 78, stated that he was unable to find the illustration either in the British Museum (NaturalHistory) or Berlin Museum, and apparently never saw it.Synonyms: Squalus (Scyliorhinus) waddii Blainville, 1816: 121 (variant spelling). Chiloscyllium modestum Günther,1871: 654, pl. 54. Holotype: British Museum (Natural History), skin of 52 cm (20.5 in) female. Cheloscyllium furvumRamsay, 1880: 97. Name only (nomen nudum), possibly quoted from a personal communication to him by W. Macleay,genus apparently an error for Chiloscyllium Müller and Henle, 1837. According to Ramsay (1880): “A new species, closelyallied to C. modestum Günth.”. Chiloscyllium furvum Macleay, 1881a: 364 (description); also Macleay, 1881b: 300(verbatim copy of earlier work). No type material mentioned, type locality “Port Jackson” [= Sydney Harbour]. Chiloscylliumfuscum Parker and Haswell, 1897: 135. No type material, Australia, possible error for C. furvum?Other Combinations: Brachaelurus modestum (Günther, 1871).FAO Names: En - Blind shark; Fr - Requin aveugle des roches; Sp - Tiburón ciego de roca.UNDERSIDE OF HEADDORSAL VIEW OF HEADField Marks: A small stout shark with long tapering barbels, nasoral grooves and circumnarial grooves, very large spiracles,a short mouth ahead of the eyes, a median symphysial groove on the chin, no dermal lobes on sides of head, twoequal-sized spineless dorsal fins and an anal fin, the first dorsal-fin origin over the pelvic-fin bases, a short precaudal tail andshort caudal fin, and colour blackish to light brown above with or without darker saddles, light yellowish on underside, usuallywith many small white spots.Diagnostic Features: See genus Brachaelurus above.Distribution: Confined to the western South Pacific offeastern Australia (southern Queensland and New SouthWales from Moreton Bay near Brisbane south to JervisBay). Records from Western Australia and northernTerritory need confirmation.Habitat: An inshore bottom shark of temperate Australianwaters, often close inshore in tidepools that are barelydeep enough to cover it and at the surf line but occasionallydown to about 73 m and exceptionally to about 137 m. Itfavours rocky shoreline areas, patches of seaweed andcoral reefs. Adults occur in caves and under ledges duringthe day, while juveniles may be in shallow areas with wavesurge in crevices and under ledges.Fig. 107 Brachaelurus waddi

146 FAO Species Catalogue for Fishery Purposes No. 1Biology: A common sluggish shark that is night-active but will take angler’s baits during the daytime. Developmentovoviviparous, with 7 or 8 young in a litter. Said to breed in summer (November in the Sydney area). Feeds on small reefinvertebrates, including crabs, shrimp, cuttlefish, squid and sea anemones, and small fishes; a coralline alga was found inthe stomach of one shark. Termed ‘blind shark’ by anglers because it retracts its eyeballs, which causes its thick eyelids toclose, when removed from the water. It can apparently live a long time out of water.Size: Maximum exceptionally to between 90 and 122 cm, most individuals smaller; an adult male was 62 cm long and anadult female, 66 cm; size at birth 15 to 18 cm.Interest to Fisheries and Human Impact: A harmless and hardy shark that thrives in aquaria. Taken offshore in bottomtrawls but not utilized commercially. Commonly caught by sports anglers with rod-and-reel from shore in rocky areas, offreefs, and in seaweed patches, particularly around Sydney and in southern Queensland. Regarded as a pest by anglers,because it sucks in baited hooks which are hard to remove from its pharynx through its small mouth and strong jaws. It maynip people when provoked. Its flesh is regarded as unpalatable because of an ammoniacal taste that is not readily removedby soaking in seawater. The conservation status of this shark is uncertain.Local Names: Brown cat-shark, Catshark or Cat shark, Dusky dogfish.Remarks: Whitley (1934: 182) suggested that Squalus waddi is the earliest name for the Australian ‘blind shark’, whichBloch and Schneider described from an illustration of an Australian shark by Dr John Latham. Although there are problemswith this interpretation (see species name above) Whitley’s substitution of waddi for modestum (which had universal useprior to Whitley’s note) for this species has been widely followed by subsequent authors.Literature: Waite (1901); Whitley (1940); Fowler (1941, 1967a); Stead (1963); Grant (1972, 1982); Shiino (1976);Compagno (1984); Dingerkus (1986); Last and Stevens (1994).Heteroscyllium Regan, 1908Genus: Heteroscyllium Regan, 1908c, Ann. Mag. Nat. Hist., (8), 2(11): 455.Type Species: Brachaelurus colcloughi Ogilby, 1907, by original designation. Replacement name for genusBrachaelurus Ogilby, 1908.Number of Recognized Species: 1.Synonyms: Genus Brachaelurus Ogilby, 1908: 2, 3. Type species: Brachaelurus colcloughi Ogilby, 1908, by originaldesignation. A junior homonym of genus Brachaelurus Ogilby, 1907.Field Marks: See Heteroscyllium colcloughi below.Diagnostic Features: Head long and slightly flattened in adults, about 22% of total length and with maximum head height0.7 to 0.8 times head width; head narrowly arched and parabolic in dorsoventral view. Snout moderately long, prenarialsnout about 2.5% and preoral snout about 5.4 to 6.0% of total length; snout wedge-shaped in lateral view, with ventralsurface of prenarial snout nearly horizontal. Eyes large and elongated-fusiform with length 2.4 to 2.5% of total length inadults; eyes not elevated above level of head. Spiracle rounded, lateral and vertical on head, just behind eyes. Nostrilsventral and well behind snout tip; nasal barbel with an expanded, hooked or rounded posterior flap at its midlength.Anal-caudal space elongated, about equal to anal-fin inner margin. Denticles small and smooth. First dorsal-fin originusually anterior to middle of pelvic-fin bases; apex anterior to insertion; free rear tip acutely angular. Second dorsal finnoticeably smaller than first and with height and base length less than 0.9 times first dorsal fin. Second dorsal-fin apexanterior to insertion; free rear tip acutely angular. Anal-fin origin about under midbase to second third of second dorsal fin;free rear tip just reaches lower caudal origin. Total vertebral count 117 to 120, precaudal count 69 to 73, monospondylousprecaudal count 30 to 35, diplospondylous precaudal count 36 to 39. Colour: background colour of the dorsal surface lightgrey-brown, sometimes golden brown, without white spots on fins and body; adults plain with indistinct saddles and whiteblotches on anterior webs of dorsal fins; young have conspicuous black blotches on posterior dorsal-fin webs and alongbase of caudal fin and dark saddles with broad light spaces between them.click for next page

click for previous pageSharks of the World, Vol. 2 147Heteroscyllium colcloughi (Ogilby, 1908) Fig. 108Brachaelurus colcloughi Ogilby, 1908, Proc. Roy. Soc. Queensland, 1907, 21: 4. “Type”: Amateur Fisherman’s Associationof Queensland Museum AFAQ no. 410, an immature male 460 mm TL (extended) according to the original description, or457 mm TL (extended) according to Ogilby and McCulloch (1908, J. Proc. Roy. Soc. N.S. Wales, 42: 285), from Mud Island,Moreton Bay, Queensland, Australia. According to Ogilby (1916, Mem. Queensland Mus. 5: 76) this specimen was“accidentally destroyed”. J. Johnson (pers. comm.), Queensland Museum, July 31, 1996) noted that the QueenslandMuseum had made an effort to locate the AFAQ specimens and failed. Ogilby (1908) also noted: “There is a secondspecimen of about the same size in the State Museum” [Queensland Museum, where J.D. Ogilby was based]. Ogilby andMcCulloch (1908) also cite this as: “A second specimen, also a young male of similar size, has been for some years in theQueensland Museum.” The description fits QM I-965, a 516 mm adolescent male from Moreton Bay which was examinedand measured by the writer and which is labelled “lectotype” in Ogilby’s handwriting (J. Johnson, pers. comm.). The twospecimens apparently are syntypes because they were both mentioned in the original description. Following Ogilby’sapparent wishes on its label, the syntype specimen QM I-965 is designated here as lectotype.Synonyms: None.Other Combinations: None.FAO Names: En - Bluegray carpet shark; Fr - Requin aveugle gris-bleu; Sp - Tiburón ciego gris.JUVENILEFig. 108 Heteroscyllium colcloughiField Marks: A small stout to slender shark with a pair of long barbels each bearing an expanded, hooked or roundedposterior flap at its midlength, nostrils with nasoral grooves and circumnarial grooves, a short mouth ahead of the eyes,symphysial groove present on the chin, no dermal lobes on sides of head, large spiracles; two spineless dorsal fins and ananal fin, the first dorsal fin larger than the second and with origin over the pelvic-fin bases, a short precaudal tail and caudalfin, and colour greyish above and white below without light spots. Young with conspicuous black and white markings onback, dorsal fins and caudal fin, fading with growth and inconspicuous in adults.Diagnostic Features: See genus Heteroscyllium above.Distribution: Apparently confined to the western SouthPacific, off Australia (southern and northeastQueensland). Most records are from Moreton Bay justeast of Brisbane, with a few records north or south of itbetween Gladstone and Coolangatta, and innortheastern Queensland off the York Peninsula and theGreat Barrier Reef.Habitat: A little-known tropical or subtropical, inshorebottom shark of the Queensland continental shelf, mostrecords from shallower than 6 m depth on soft bottom oraround wrecks.UNDERSIDE OF HEADBiology: A rare to uncommon species, with biologypoorly known. Presumably feeds on benthicinvertebrates, but diet not recorded. Ovoviviparous(aplacental viviparous), with litters of six or seven young. Apparently egg-cases are lost early in development, and term ornear-term foetuses are not enclosed in hard or soft egg-cases. Near-term foetuses 164 to 168 mm long had relatively large

148 FAO Species Catalogue for Fishery Purposes No. 1yolk-sacks filled with yolk but these were lost in term foetuses 174 to 186 mm long, suggesting that foetal nutriment isprimarily through vitelline yolk. As with Brachaelurus waddi this species also closes its eyes when removed from the water(J. Johnson, pers. comm.).Size: Maximum to at least 75.5 cm, pregnant females 65.8 to 75.5 cm, males adolescent at ca. 48.2 to 51.6 cm. Termfoetuses 174 to 184 mm, size at birth probably between 17 to18 cm.Interest to Fisheries and Human Impact: A harmless shark of no interest to fisheries but occasionally caught by sportsanglers. It is partly sympatric with Brachaelurus waddi off southern Queensland and has been confused with it in the past.By analogy to B. waddi and because of its inshore benthic habitat this is likely to be a hardy aquarium shark and an attractiveexhibit (particularly the boldly barred young). Some of these sharks have been kept in aquaria (J. Stevens, pers. comm.). Ifthis species enters the aquarium trade in any numbers this should be strictly regulated.Restricted habitat, distribution in areas heavily and increasingly utilized by people, and possibly rarity suggested by fewrecords in suitable habitat in the relatively well-surveyed waters where it occurs makes it inherently vulnerable to depletion.Its distribution, behaviour, ecology, and conservation status need to be studied in detail by diving surveys. Surveys of anglercatches and commercial demersal landings need to be made to determine its presence in inshore fisheries. This specieshas a status of Vulnerable in the IUCN 2000 Red List.Local Names: Colclough’s shark or Colcloughs shark, Blue-grey catshark, Blue-grey shark.Literature: Ogilby (1908, 1916); Ogilby and McCulloch (1908); Garman (1913); McCulloch and Whitley (1925); Whitley(1934, 1940); Fowler (1929, 1941, 1967a); White (1937); Bigelow and Schroeder (1948); Grant (1972, 1982); Compagno(1973, 1981b, 1982, 1984); Applegate (1974); Dingerkus (1986); Last and Stevens (1994); J. Johnson (pers. comm.), P.Last (pers. comm.), J. Stevens (pers. comm.).2.3.3 Family ORECTOLOBIDAEFamily: Orectolobidae Gill, 1896, Proc. U.S. Natl. Mus. 18(1057): 212. Also subfamily Orectolobinae Fowler, 1934, Proc.Acad. Nat. Sci. Philadelphia, 85: 238 (Family Orectolobidae).Type Genus: Orectolobus Bonaparte, 1834.Number or Recognized Genera: 3.Synonyms: Subfamily Crossorhinae Swainson, 1839: 318 (Family Squalidae). Type genus: Crossorhinus Müller andHenle, 1837.FAO Names: En - Wobbegongs; Fr - Requins-tapis; Sp - Tiburones tapiceros.Field Marks: These are distinctive flattened, variegated sharks, differing from all others (except angel sharks, Squatinidae),having narrow dermal flaps along the sides of the head. Wobbegongs differ from angel sharks in having anal fins andseparate cermal lobes among many other differences. Wobbegongs also have long barbels, short, nearly terminal mouths infront of the eyes, nasoral grooves and circumnarial grooves and flaps, symphysial grooves, large spiracles and dorsolateraleyes.Diagnostic Features: Head very broad and flattened, with unique lateral flaps of skin. Snout truncated. Eyes dorsolaterallysituated on head and with strong subocular ridges below them. Eyes without movable upper eyelids but with subocularpockets and ridges. Spiracles very large and larger than eyes, with prominent raised external rims; spiracles somewhatbelow, behind and lateral to eyes. Gill slits small, fifth gill slit well-separated or close to fourth but not overlapping it; internalgill slits without filter screens. Nostrils with very long pointed or branched barbels, circumnarial folds and circumnarialgrooves around outer edges of incurrent apertures. Nasoral grooves short and strongly developed. Mouth large, slightlyarched and nearly transverse, and nearly terminal on head. Lower lip not trilobate and without lateral orolabial groovesconnecting edge of lip with medial ends of lower labial furrows, but with a longitudinal symphysial groove on chin. Lowerlabial furrows extending medially nearly to symphysis, but not connected medially by a mental groove or groove and flap.Teeth strongly differentiated in upper and lower jaws, with three rows of fang-like teeth at the upper symphysis and two rowsat the lower. Tooth row count 23 to 26/19. Teeth with a strong medial cusp, lateral cusplets variably present or absent, andlabial root lobes weak. Teeth orthodont with a central pulp cavity and no plug of osteodentine. Body considerably depressed,without ridges on sides. Precaudal tail shorter than body. Caudal peduncle without lateral keels or precaudal pits. Pectoralfins moderate-sized or very large, broad and rounded. Pectoral fins aplesodic and with fin radials not expanded into fin web.Pectoral propterygium large and separate from mesopterygium and metapterygium; pectoral-fin radial segments three atmost, and with longest distal segments about 0.5 times the length of longest proximal segments. Pelvic fins larger thandorsal and anal fins, nearly as large as pectoral fins and with anterior margins 0.6 to 0.8 times the pectoral-fin anteriormargins. Claspers without mesospurs, claws or dactyls. Dorsal fins equal-sized, first dorsal-fin origin over or slightly behind

Sharks of the World, Vol. 2 149the pelvic-fin insertion, insertion far behind the pelvic-fin rear tips. Anal fin about half as large as second dorsal fin or less,with broad base, subangular or narrowly rounded apex, origin about opposite rear third of second dorsal-fin base or itsinsertion, and insertion separated by a narrow notch much less than base length from lower caudal-fin origin. Caudal finshort and not crescentic, weakly heterocercal with its upper lobe at a low angle above the body axis; dorsal caudal-fin marginless than a fourth as long as the entire shark. Caudal fin with a strong terminal lobe and subterminal notch but without aventral lobe, preventral and postventral margins not differentiated and forming a continuous curve. Vertebral centra withwell-developed radii. Total vertebral count 149 to 158, monospondylous precaudal count 41 to 52, diplospondylousprecaudal count 42 to 62, diplospondylous caudal count 49 to 62, and precaudal count 87 to 106. Cranium moderately broadand not greatly expanded laterally. Medial rostral cartilage extremely short and reduced to a low projection. Nasal capsulesgreatly depressed and with double longitudinal fenestrae, internarial septum compressed but low, not expanded laterally.Orbits with small foramina for preorbital canals, medial walls not fenestrated around the optic nerve foramina. Supraorbitalcrests present on cranium but not laterally expanded and pedicellate. Suborbital shelves very broad and not reduced.Cranial roof without fenestrae. Basal plate of cranium with separate pairs of stapedial and carotid foramina. Adductormandibulae muscle of jaws with two divisions. Preorbitalis muscles not extending onto posterodorsal surface of cranium. Noanterodorsal palpebral depressor, rostromandibular, rostronuchal or ethmonuchal muscles. Valvular intestine of ring typewith 23 to 33 turns. Development ovoviviparous. Size small to large with adults between 60 cm and at least 3 m. Colourpattern highly developed, including dark and light spots, dark saddles, rings and reticulations on back.Distribution: Wobbegongs are common, largish flattened bottom sharks that are endemic at present to warm-temperate totropical continental waters of the western Pacific. They are most diverse in Australian and New Guinean waters, but occurnorthwards to Japan.Habitat: Wobbegongs occur in inshore and offshore bottom habitats from the intertidal down to at least 110 m. They areoften found on rocky and coral reefs or on sandy bottom, where they lurk and are concealed in part by their cryptic colorationand dermal lobes on their heads.Biology: Wobbegongs are reportedly sluggish fishes, moving little, but can clamber around with their paired fins on thebottom and even climb partway out of the water while moving between tidepools. At least two species may attain a size of 3.2to 3.7 m. These sharks are ovoviviparous, with large litters of 20 or more young. Wobbegongs are powerful bottom predatorswith heavy jaws and greatly enlarged, dagger-shaped, extremely sharp teeth in the front of their mouths. They feed onbottom fishes, crabs, lobsters, octopi, and other bottom animals.Interest to Fisheries and Human Impact: Wobbegongs are utilized for food in Australia and off China, Japan, Malaysia(Sabah) and probably elsewhere where they occur; their colourful skins are also used for leather. Wobbegongs can bitewhen captured or when provoked or stepped upon, and have inflicted severe lacerations on the limbs of people. There arerare records of large wobbegongs biting off people’s feet or killing them, but these need to be verified. Wobbegongs are oftendifficult to see against the bottom and can be contacted accidentally. These sharks should be treated with respect becauseof their strong dentition, as with angel sharks (Squatinidae), even though they do not appear to be particularly prone to bitepeople unless provoked. Michael (1993) thought that these sharks had poor visual acuity and tended to bite at any objectthat moved near their heads, so that people should avoid placing their limbs near them. Wobbegongs are regularly kept inpublic aquaria in Australia, Europe, and the United States, and are often viewed by divers on the Great Barrier Reef ofAustralia and probably off Japan. They have bred in captivity.Local Names: Wobbegongs, Carpetsharks, Carpet sharks; Kovrovye akuly (Russian).Remarks: The arrangement of this family follows Compagno (1984), Dingerkus (1986), and Last and Stevens (1994).Although apparently a monophyletic group and the sister taxon of the Brachaeluridae (Dingerkus, 1986; Compagno, 1988),the Orectolobidae requires a detailed morphological and genetic investigation to elucidate the interrelationships of itscomponent taxa. Dingerkus (1986) suggested that the highly derived Eucrossorhinus was the sister genus of Orectolobus,and both formed the sister taxon of Sutorectus, but this is complicated by Orectolobus sp. A, which has some characters,including its narrow interdorsal space, slightly lower dorsal fins, and longitudinal rows of small dermal knobs and ridges inyoung, that suggest that it is intermediate between typical Orectolobus and the bizarre Sutorectus tentaculatus.Literature: Ogilby and McCulloch (1908); Regan (1908a); Garman (1913); Whitley (1940); Fowler (1941); Compagno(1984); Dingerkus (1986); Michael (1993); Last and Stevens (1994).Key to Genera:1a. Chin with highly branched dermal lobes; dermallobes on sides of head mostly extensivelybranched (Fig. 109); body with a reticular patternof narrow dark lines . . . . . . . . . . . . Eucrossorhinus1b. Chin without dermal lobes; dermal lobes on sidesof head mostly simple or with a few branches(Fig. 110); colour pattern variable, but without areticular pattern of narrow dark lines . . . . . . . . . . 2dermal lobeson chinUNDERSIDE OF HEADFig. 109 Eucrossorhinusno dermallobes on chinUNDERSIDE OF HEADFig. 110 Orectolobus

150 FAO Species Catalogue for Fishery Purposes No. 12a. Head and body without tubercles or withsmall ones or inconspicuous dermalridges in young (lost in adults); dorsal finshigh, height at least three-fourths of baselength; origin of first dorsal fin behindmidbases of pelvic fins (Fig. 111). . . . . . . Orectolobus2b. Head and body covered by large roundedtubercles; dorsal fins long and low, heightabout half of base length; origin of firstdorsal fin anterior to pelvic-fin midbases(Fig. 112) . . . . . . . . . . . . . . . . . . . Sutorectusdorsal fins highFig. 111 Orectolobustuberclesdorsal fins lowFig. 112 SutorectusEucrossorhinus Regan, 1908Genus: Eucrossorhinus Regan, 1908a, Proc. Zool. Soc. London, 1908: 357.Type Species: Eucrossorhinus dasypogon Regan, 1908, by monotypy, equals Crossorhinus dasypogon Bleeker, 1867.Number of Recognized Species: 1.Synonyms: Genus Crossohrinus Compagno, 1984: 179. Typographical error for Crossorhinus dasypogon Bleeker, 1867.Diagnostic Features: Head very broad, its width slightly greater than its length from snout tip to fifth gill openings. Chin witha bushy beard of highly branched dermal lobes. Dermal lobes of sides and front of head highly branched and numerous, inapproximately 24 to 26 pairs, forming a virtually continuous fringe from snout tip to pectoral-fin bases. Nasal barbelsbranched, with complex multiple lobes. Mouth broad, width about 11% of total length. Head and body without enlargedtubercles on body, except for those above eyes. Trunk very broad, width across pectoral-fin insertions about equal to headlength. Precaudal tail rather short, distance from pelvic-fin insertion to lower caudal origin about equal to head length.Interspace between first and second dorsal fins longer than first dorsal-fin inner margin and slightly more than half firstdorsal-fin base. Pectoral and pelvic fins very large, distance from pectoral-fin insertions to pelvic-fin origins about equal topectoral-fin bases and less than pelvic-fin lengths from origins to free rear tips. Dorsal fins high and short, height of firstdorsal fin about equal to its base length, length of first dorsal fin base less than pelvic-fin length. Origin of first dorsal finopposite posterior fourth of pelvic-fin bases. Colour: dorsal surface with a reticular pattern of narrow dark lines on a lightbackground, with scattered symmetrical enlarged dark dots at the junction of lines.Remarks: This genus was originally proposed by Regan (1908a) to separate Crossorhinus dasypogon Bleeker, 1867 fromother wobbegongs primarily because of its supposedly even-spaced gill slits, but he also mentioned that the genus differedfrom Orectolobus by having a broader, more depressed head, smaller eyes and wider spiracles. However, Regan (1909)reversed himself and rejected Eucrossorhinus because his new, very similar (and ultimately conspecific) Orectolobusogilbyi had the last two gill slits close together.Ogilby and McCulloch (1908), Fowler (1941), and Stead (1963) did not recognize the genus Eucrossorhinus but Garman(1913), Whitley (1940), Bigelow and Schroeder (1948), Garrick and Schultz (1963), Compagno (1973, 1984), Applegate(1974), Dingerkus (1986), and Last and Stevens (1994) all retained it. Eucrossorhinus is morphologically divergent fromother wobbegongs, but its phyletic relationships are unclear and require a detailed study of its morphology.Two species of Eucrossorhinus have been recognized, E. dasypogon from Indonesia, and E. ogilbyi from Torres Straitsand Papua-New Guinea. Regan (1909) distinguished the two as follows:[Eucrossorhinus] ogilbyi: Gill slits decreasing in size from first to fourth, last larger; last two closer together thanrest. Dermal lobes on sides of head in three separate groups. Origin of first dorsal fin well behind middle of totallength. Distance between origins of dorsals nearly half that from origin of second dorsal fin to end of tail.[Eucrossorhinus] dasypogon: First gill slit slightly smaller than rest, which are of equal size and equidistant. Dermallobes on head in two separate groups. Origin of first dorsal fin in the middle of total length. Distance between originsof dorsals slightly more than one-third that from origin of second dorsal-fin to end of tail.Fowler (1941) used Eucrossorhinus as a subgenus of Orectolobus, but placed O. dasypogon in Eucrossorhinus andOrectolobus ogilbyi in the subgenus Orectolobus. Compagno (1984) compared a possible syntype of Eucrossorhinusdasypogon collected by P. Bleeker from Indonesia (see below) with a larger specimen labelled E. ogilbyi from northern

Sharks of the World, Vol. 2 151Queensland (BMNH 1911.4.1.43, 415 mm female), as well as a much larger specimen labelled E. dasypogon from NewGuinea (Australian Museum, Sydney, AMS I4783, 117 cm adult male). This suggested that the characters supposed toseparate the two species did not hold. All specimens had the last two gill openings more closely spaced than the first three,though the New Guinea and Queensland specimens had them slightly closer than the small Indonesian specimen. All threespecimens have the first four gill openings about equal in length or with the first slightly smaller; the fifth is slightly smallerthan the fourth in the Indonesian specimen, slightly larger in the large New Guinea adult, and about equal to it in theQueensland specimen. In the two smaller specimens the first dorsal-fin origin is actually slightly ahead of the midlength, butslightly behind in the New Guinea adult, suggesting allometric increase in abdominal length with growth. The Indonesianspecimen has the distance from the second dorsal-fin origin to the caudal-fin tip 2.7 times the space between the origins ofthe first and second dorsal fins, the Queensland specimen 2.6 times, and the large New Guinea specimen 2.4 times.Compagno (1984) synonymized E. ogilbyi with E. dasypogon, and suggested that the differences listed between the twospecies in the literature and in the specimens he had examined represented individual and ontogenetic variation in a singlespecies. The specimens were strikingly similar in colour pattern and general morphology, and Compagno indicated thatthere was nothing in the literature to suggest any significant differences between Australian, Papua-New Guinean andIndonesian Eucrossorhinus. This was accepted by Last and Stevens (1994).Eucrossorhinus dasypogon (Bleeker, 1867) Fig. 113Crossorhinus dasypogon Bleeker, 1867, Arch. Neerl. Sci. Nat. 2: 400, pl. 21, fig. 1. Syntypes: Rikjsmuseum van NatuurlijkeHistoire, Leiden, RMNH 7411 (1) Waigiu (Waigeo), RMNH 5118 (1) Aru, Indonesia, according to Eschmeyer (1998, Cat.Fish.: CD-ROM). Uncertain status: British Museum (Natural History), BMNH-1867.11.28.209, 215 mm immature male andnoted as a syntype by Compagno (1984, FAO Fish. Syn. (125), 4 (1): 179) from documentation on the specimen.Synonyms: Orectolobus ogilbyi Regan, 1909: 529. New name for Orectolobus dasypogon Ogilby and McCulloch, 1908:272, pl. 43, fig. 1. Syntypes: Two specimens from Torres Strait and Samarai, Papua-New Guinea, including one 1 210 mmlong, according to Ogilby and McCulloch. Eschmeyer (1998: CD-ROM) identified one of these as Australian Museum,Sydney, AMS I.5405.Other Combinations: Orectolobus dasypogon (Bleeker, 1867).FAO Names: En - Tasselled wobbegong; Fr - Requin-tapis barbu; Sp - Tapicero barbudo.LATERAL VIEWUNDERSIDE OF HEADFig. 113 Eucrossorhinus dasypogon

152 FAO Species Catalogue for Fishery Purposes No. 1Field Marks: This squat, broad, angler-like shark is unmistakable, with profuse, highly branched dermal lobes on its head, abeard of similar lobes on its chin, and reticulated colour pattern of narrow dark lines and dark spots at their junctions on alight background. It is also recognizable by having its mouth in front of eyes, a symphysial groove on chin, very broadpectoral and pelvic fins, two spineless dorsal fins and an anal fin, the first dorsal-fin origin opposite the pelvic-fin hindbases,and the anal-fin origin well behind the second dorsal-fin origin.Diagnostic Features: See genus Eucrossorhinus above.Distribution: Western South Pacific: Indonesia(Waigeo, Aru), New Guinea, Malaysia?, northernAustralia (northern Queensland, Northern Territory,Western Australia).Habitat: A little-known inshore and offshore tropicalbottom shark, present inshore and on coral reefs,commonly seen on coral heads, and in reef channelsand reef faces. Michael (1993) suggested that thisspecies was an obligate coral-reef dweller.Biology: Commonly seen on the northern Great BarrierReef; rests on the bottom with its tail curled. Thisnocturnal species is said to be a faster swimmer thanother wobbegongs (Whitley and Pollard, 1980), but itsmore flattened shape, exquisite camouflage, and bushydermal flaps suggests the reverse, that it might be more sluggish than other wobbegongs. It is thought to be solitary(Michael, 1993), and individuals are seen resting in caves and under ledges in the day, but may leave their retreats at night tofeed. Individuals apparently have a small home range and several retreats within it. Probably ovoviviparous, thoughreproductive biology is little known. Feeds on bottom fishes and possibly invertebrates, and is known to eat nocturnal teleostfishes such as squirrelfish and soldierfish (Holocentridae) and sweepers (Pempheridae) that share their caves.Size: Maximum said to be 366 cm but this is uncertain and thought by Last and Stevens (1994) to be incorrect; maximumreliably to 125 cm. Born at about 20 cm; the 215 mm possible syntype in the British Museum (Natural History) is newborn orclose to it. Reaches 125 cm; an adult male from New Guinea with calcified claspers and examined by the writer (see above)was only 117 cm long.Interest to Fisheries and Human Impact: Interest to fisheries uncertain and probably minimal; the tough skin with itshandsome reticulated colour pattern is occasionally used for leather.According to an informant quoted by Whitley (1940), this shark “...attacks and generally kills the natives” in Papua-NewGuinea, but this is questionable. This shark no doubt should be treated with respect like other wobbegongs, but its fearsomereputation may be greatly exaggerated and may well evaporate as its biology becomes better known. Divers commonlyapproach and photograph the tasselled wobbegong during the daytime, without inciting agonistic behaviour, thoughprobably stepping on or near this well-camouflaged shark might cause it to bite at least in self-defence or by possiblymistaking a human foot for prey. Michael (1993) thought that it was more likely to bite people than other wobbegongs, andhas bitten divers without provocation several times. It is kept in aquaria in the United States and probably elsewhere and isviewed by ecotouristic divers in Australia. It makes a spectacular subject for display and photography.The conservation status of this species is uncertain, but should be of concern because of its limited distribution and habitaton reefs, including poorly protected areas outside Australian territorial waters that are subject to habitat destruction frompollution and bad fishing practices such as dynamiting, netting and poisoning reefs.Local Names: Bearded wobbegong, Ogilby’s wobbegong, Tasselled wobbegong.Literature: Ogilby and McCulloch (1908); Regan (1908a, c, 1909); Garman (1913); Whitley (1940); Fowler (1941); Marshall(1965); Whitley and Pollard (1980); Compagno (1984); Dingerkus (1986); Last and Stevens (1994).Orectolobus Bonaparte, 1834Genus: Subgenus Orectolobus Bonaparte, 1834, Iconog. Fauna Italica, 3, Pesci, fasc. 7. (Genus Scyllium Cuvier, 1816).Type Species: Squalus barbatus Gmelin, in Linnaeus and Gmelin, 1788, by subsequent restriction of Gill, 1896, Proc. U. S.Natl. Mus. 18(1057): 211; a junior synonym of Squalus maculatus Bonnaterre, 1788.Number of Recognized Species: 5.

Sharks of the World, Vol. 2 153Synonyms: Genus Crossorhinus Müller and Henle, 1837a: 113. Type species: Squalus lobatus Bloch and Schneider,1801, by monotypy. Genus Chrossorhinus (Müller and Henle) Smith, 1837: 86; Smith, 1838: 73. Type species: Squaluslobatus Bloch and Schneider, 1801, by monotypy. Error or emendation for Crossorhinus Müller and Henle, 1837.Diagnostic Features: Head narrow, its greatest width about equal to or less than distance from snout tip to first gillopenings. Chin smooth, without a beard of dermal lobes. Dermal lobes of sides and front of head small, short, unbranched orslightly branched, and in 3 to 10 pairs, forming isolated groups that are broadly separated from one another. Nasal barbelssimple and unbranched or with a weak basal branch. Mouth narrow, width about 9% of total length. Dorsal surface of head,body and precaudal tail, and dorsal fin bases smooth or with small inconspicuous tubercles or low longitudinal ridges, notnoticeably warty. Trunk moderately broad, width across pectoral-fin insertions considerably less than head length.Precaudal tail rather long, distance from pelvic-fin insertion to lower caudal-fin origin much greater than head length.Pectoral and pelvic fins small and widely spaced from each other, distance from pectoral-fin insertions to pelvic-fin origins atleast 1.5 times length of pectoral-fin bases and somewhat greater than pelvic-fin lengths from origins to free rear tips.Interspace between first and second dorsal fins usually slightly longer than first dorsal-fin inner margin but slightly shorter inOrectolobus sp. A, and varying from over half to about a fifth of first dorsal-fin base. Dorsal fins fairly high and short tomoderately long, height of first dorsal fin over three-fourths of its base length, length of first dorsal-fin base less thanpelvic-fin length from origin to free rear tip. Origin of first dorsal fin behind midbases of pelvic fins. Dorsal surface with acolour pattern of regular or jagged-edged broad dark saddles separated by light areas with dusky blotches, scattered darkspots or semi-reticulated broad lines, or O-shaped light spots on a dark background; no reticulating narrow lines with spotsat their junctions, but broad reticulating lines without spots are present in a few species.Remarks: The present account of this genus follows Ogilby and McCulloch (1908), Regan (1908a), Whitley (1940),Compagno (1984), and Last and Stevens (1994). As with Compagno (1984), the account is regarded as provisional, withsome problems including the status of subspecies in Orectolobus ornatus. Also, in Western Australian waters there is adistinct, undescribed species of wobbegong, that is very abundant and resembles O. ornatus as well as Sutorectustentaculatus (B. Hutchins, pers. comm.). It is termed Orectolobus sp. A by Last and Stevens (1994) and in the presentaccount, and is being described by P. Last, B. Hutchins, and the writer. There is also a record of a member of this genus fromSabah, Borneo (Fowler et al., 1999), based on a dried specimen of Orectolobus that is of uncertain placement in the genus.A wobbegong from Japan illustrated by Nakaya and Shirai (1984, pl. 8, fig. B) is identified as O. ornatus but may be anundescribed species. Its coloration of broad black rings and reticulations on a yellowish background and reduced saddles isvery different from the Australian O. ornatus and unlike any other wobbegong.Key to Species:1a. Nasal barbels not branched (Fig. 114);dermal lobes of head very broad-based,only 2 or 3 in front of eyes; colour patternsimple, with a few dark spots, duskymottling, and three large dark, light-edgedrounded saddles on back anterior to firstdorsal fin . . . . . . . . . . Orectolobus wardi1b. Nasal barbels branched (Fig. 115); dermallobes narrower-based and morenumerous, four or more in front of eyes;colour pattern with elaborate variegatedspots and saddles . . . . . . . . . . . . . 2simple barbelsUNDERSIDE OF HEADFig. 114 Orectolobus wardi6-10lobesbranched barbelsUNDERSIDE OF HEADFig. 115 Orectolobus maculatus2a. About 6 to 10 dermal lobes below and infront of eyes; back dark, with whiteO-shaped spots and white blotchesobscuring darker saddles (Figs 115 and116) . . . . . . . . . . Orectolobus maculatus2b. About 4 to 6 dermal lobes below and infront of eyes; back with dark colourvariegated with light blotches andprominent saddle markings . . . . . . . . 3Fig. 116 Orectolobus maculatus

154 FAO Species Catalogue for Fishery Purposes No. 13a. About four dermal lobes below and infront of eyes; dermal lobes behindspiracle unbranched and slender;interdorsal space slightly shorter thaninner margin of first dorsal fin (Fig. 117). . . . . . . . . . . . . . . . . Orectolobus sp. A 4 dermallobes5-6 dermallobesbroadlobesbehindspiracle3b. About 5 or 6 dermal lobes below and infront of eyes; dermal lobes behindspiracle branched, or if unbranched verybroad; interdorsal space slightly longerthan inner margin of first dorsal finUNDERSIDE OF HEADFig. 117 Orectolobus sp. A(Fig. 118). . . . . . . . . . . . . . . . . . . . 4Fig. 118 Orectolobus ornatussaddles not black edgedUNDERSIDE OF HEAD4a. Back with light areas between darksaddles marked with broad reticulateddark lines; borders of saddles dark but notconspicuously black-edged (Fig. 119). . . . . . . . . . . . . . . Orectolobus japonicus4b. Back with light areas between darksaddles marked with dark, light centredblotches and spots, not reticulated lines;saddles with conspicuous black borders(Fig. 120) . . . . . . . . . . . Orectolobus ornatusFig. 119 Orectolobus japonicusblack-edged saddlesFig. 120 Orectolobus ornatusOrectolobus japonicus Regan, 1906 Fig. 121Orectolobus japonicus Regan, 1906b, Ann. Mag. Nat. Hist. (7), 18(108): 435. Syntypes: British Museum (Natural History),BMNH 1862.11.1.18 and BMNH 1862.11.1.90 according to Eschmeyer (1998, Cat. Fish.: CD-ROM), two females, 1 000 and780 mm TL, Japan.Synonyms: None.Other Combinations: Crossorhinus or Orectolobus barbatus (not Gmelin, 1788), Crossorhinus lobatus (not Bloch andSchneider, 1801).FAO Names: En - Japanese wobbegong; Fr - Requin-tapis moustache; Sp - Tapicero japonés.Fig. 121 Orectolobus japonicus

Sharks of the World, Vol. 2 155Field Marks: Flattened benthic sharks with dermal lobes on sides of head, symphysial groove on chin, very conspicuousvariegated colour pattern of broad dark dorsal saddles with light spots and corrugated edges, interspaced with light areaswith dark broad reticular lines; also, mouth in front of eyes, long, basally branched nasal barbels, nasoral grooves andcircumnarial grooves, two rows of enlarged fang-like teeth in upper jaw and three in lower jaw; first dorsal-fin origin overpelvic-fin bases.Diagnostic Features: Nasal barbels with a few branches. Five dermal lobes below and in front of eye on each side of head;dermal lobes behind spiracles branched and broad. No dermal tubercles or ridges on back. Interspace between dorsal finslonger than inner margin of first dorsal fin, about half first dorsal-fin base. Origin of first dorsal fin behind midbases of pelvicfins. First dorsal-fin height about equal to base length. Colour: colour pattern highly variegated and conspicuous, dorsalsurface of body with conspicuous broad, dark rectangular saddles with deeply corrugated margins, dotted with light spotsand not ocellate in appearance; interspaces between saddles light, with numerous broad reticulated lines.Distribution: Western North Pacific: off Japan, Korea,China, Taiwan (Province of China), Viet Nam, andPhilippines.Habitat: A little-known temperate to tropical inshore bottomshark, found on rocky and coral reefs.Biology: A nocturnal shark, rarely observed by divers.Ovoviviparous, with litters of up to 20 to 23 young. In captivityin an aquarium in Okinawa, Japan, gives birth in spring(March through May), and has been observed in courtship.The male grabs the female in the gill area while mating withher. The gestation period is about a year. Eats primarilybenthic fish, including lizardfishes (Synodontidae),cutlassfish (Trichiuridae), horse mackerel and other jacks(Carangidae), goatfishes (Mullidae), groupers (Serranidae),tilefishes (Malacanthidae), sea robins (Triglidae), whiting(Sillaginidae), parrotfishes (Scaridae), sea bream(Sparidae), croakers (Sciaenidae), also skates (Rajidae),shark egg-cases, cephalopods, and shrimp.Size: Maximum uncertain, reaches at least 107 cm. Size atbirth 21 to 23 cm; a male was mature at 103 cm; adultfemales that gave birth in captivity were 101 to 107 cm.Interest to Fisheries and Human Impact: Interest to fisheries probably limited, caught in set nets in Japan and used forhuman consumption; also taken in China, Taiwan (Province of China), Korea and Viet Nam. Conservation status uncertain.Probably viewed by ecotouristic divers in Japan, and kept in aquaria in Japan and the United States.Local Names: Japanese carpet shark, Carpet shark, Bearded shark, Japanese bearded shark, Fringe shark, Kirinotobuka,Oose or Ôse (Japan).Literature: Ogilby and McCulloch (1908); Regan (1908a); Garman (1913); Fowler (1941); Herre (1953); Lindberg andLegeza (1959); Fourmanoir and Nhu-Nhung (1965); Matsubara (1955); Chen (1963); Masuda, Araga and Yoshino (1975);Uchida (1982); Compagno (1984); Nakaya and Shirai (1984); Uchida, Toda and Kamei (1990); Michael (1993).Orectolobus maculatus (Bonnaterre, 1788) Fig. 122Squalus maculatus Bonnaterre, 1788, Tabl. Encyclop. Method. Trois Reg. Nat., Ichthyol., Paris: 8. No type material? Typelocality: “La mer du sud”.Synonyms: Squalus barbatus Gmelin, in Linnaeus and Gmelin, 1788: 1493. New Holland. On ‘barbu’ of Broussonet, 1780,Mem. Acad. Sci. Paris, 657, no. 7, New Holland. No types known according to Eschmeyer (1998: CD-ROM). Squaluslobatus Bloch and Schneider, 1801: 137. On the ‘Watt’s Shark’ of Phillip, 1789: 285, pl. 53. Sydney Cove, Port Jackson, NewSouth Wales, Australia. No types known according to Eschmeyer (1998: CD-ROM). Squalus appendiculatus Shaw andNodder, 1806: pl. 737. Antarctic seas. On the ‘Watt’s Shark’ of Phillip, 1789: 285, pl. 53. Sydney Cove, Port Jackson, NewSouth Wales, Australia. No types known according to Eschmeyer (1998: CD-ROM).Other Combinations: Crossorhinus, Orectolobus, Scyllium, Chiloscyllium, orScyliorhinus barbatus (Gmelin, 1788),Chrossorhinus or Crossorhinus lobatus (Bloch and Schneider, 1801).

156 FAO Species Catalogue for Fishery Purposes No. 1FAO Names: En - Spotted wobbegong; Fr - Requin-tapis tacheté; Sp - Tapicero manchado.DORSAL VIEWLATERAL VIEWUNDERSIDE OF HEADField Marks: Flattened benthic sharks with dermal lobes on sides of head, symphysial groove on chin; a rather sombre,variegated colour pattern of dark back with obscure darker dorsal saddles and densely covered with prominent lightO-shaped spots; also, mouth in front of eyes, long, basally branched nasal barbels, nasoral grooves and circumnarialgrooves, two rows of enlarged fang-like teeth in upper jaw and three in lower jaw; first dorsal-fin origin over pelvic-fin bases.Diagnostic Features: Nasal barbels with a few basal branches. Six to ten dermal lobes below and in front of eye on eachside of head; dermal lobes behind spiracles branched and broad. No dermal tubercles or ridges on back. Interspacebetween dorsal fins longer than inner margin of first dorsal fin, about half length of dorsal-fin base. Origin of first dorsal finover about last third of pelvic-fin base. First dorsal-fin height about equal to base length. Colour: colour pattern variegatedbut more sombre and less contrasting than most other wobbegongs except O. wardi, dorsal surface of body dark withsomewhat obscure, broad, darker rectangular saddleswith deeply corrugated margins separated by lighterareas, the entire dorsal surface densely spotted withlarge, O-shaped, light markings; saddles not ocellate inappearance; interspaces between saddles without broadreticulated lines.Distribution: Western Pacific: South coast of Australia(Western Australia, South Australia, Victoria, New SouthWales and southern Queensland, Tasmanian recordsprobably invalid, possibly not Northern Territory). Recordsfrom Japan and the South China Sea need confirmation.Habitat: An abundant, temperate to tropical, inshore tooffshore bottom shark of the continental shelves of thewestern Pacific, occurring in the intertidal down to at least110 m, commonly on coral and rocky reefs, in coastalbays, in estuaries, in seagrass beds, under piers, and onsandy bottom. It may occur in water barely deep enoughto cover it, and has been seen climbing over ridgesbetween tidepools, with its back out of water. Juvenilesoccur on low reefs, in seagrass beds, and in estuaries. Itsometimes makes short trips well above the bottom.Biology: This shark apparently is sluggish and inactiveand is often found motionless on the bottom, at leastduring the day when it is presumably resting. It often isfound in caves, under overhangs on rocky reefs, inchannels, and in shipwrecks during the day. It is wellcamouflaged by its colour pattern and dermal flaps onFig. 122 Orectolobus maculatus

Sharks of the World, Vol. 2 157rough bottom but is rather conspicuous on sand. This species (and wobbegongs in general) has not been studied to theextent of some nurse sharks (Ginglymostomatidae), but site specificity may be a feature of its behaviour as with nursesharks: anecdotal accounts suggest that individuals may return to the same site repeatedly. It may occur singly but alsooccurs in aggregations of a dozen or more. It is said to be nocturnal, and may swim and clamber about the bottom at nightlooking for food as nurse sharks do. It is not known how important their camouflage patterns are for feeding in this and otherwobbegongs. It is uncertain if wobbegongs take a substantial amount of prey that simply blunders into proximity while theysit on the bottom (as shown in a recent video), or if they do so by active prowling and stalking at night. Wobbegongs in theSydney area, presumably this common species, were observed to slowly sneak up to a bait at night from a considerabledistance, as if stalking potential prey like a cat, but this may not be the case with live, uninjured prey.Ovoviviparous, with large numbers of young per litter; one female had 37. There are anecdotal accounts that malewobbegongs from the Sydney area (and presumably this species, which is abundant there) kept in aquaria fight vigorouslyamong themselves while courting females, and that females are bitten by males in the gill region during courtship and oneclasper is inserted; in captivity, these wobbegongs copulated in July. A wild male wobbegong was said to be attracted to afemale kept in a wired enclosure open to the sea and tried to enter the enclosure during the breeding season; the implicationis that the female gave off an attractive stimulus, presumably a chemical pheromone but possibly some other signal.The spotted wobbegong feeds on bottom invertebrates, including crabs, lobsters and octopuses, bony fishes including seabass (Serranidae), scorpionfishes (Scorpaenidae) and luderick (Kyphosidae), other nonbatoid sharks includingconspecifics, and rays (batoids). Prey items may blunder right up to the mouth of a lurking wobbegong, and even nibble onits tentacles, before being caught and eaten. Apparently the short broad mouth and large broad pharynx of this and otherwobbegongs aid them in sucking in prey. Video footage suggests that prey is suddenly sucked into the mouth as the pharynxexpands, much as in angel sharks, but the prey is taken in front of the wobbegong rather than above it as in angel sharks(Squatinidae). The powerful jaws and big, modified anterior teeth in the symphysial region of this and other wobbegongs,with one median and two lateral rows of teeth in the lower jaw that interdigitate with two rows of enlarged lateral teeth in theupper jaw, form an effective trap to impale and kill their prey.Size: Maximum about 320 cm, but with most individuals smaller, up to 150 to 180 cm. Size at birth about 21 cm. Adult malesmay mature at about 60 cm long.Interest to Fisheries and Human Impact: Interest to fisheries limited, utilized for human consumption and for leather; themeat is apparently excellent eating and the skin of this and other wobbegongs is tough and makes an excellent, decorativeleather with its handsome patterning. Spotted wobbegongs are commonly caught as bycatch in trawls, beach seines,trammel nets, in lobster pots and traps, and are fished with line gear (droplines) off New South Wales. Some are taken bydivers with spears. These sharks are regarded as a pest by lobster fishers, because they are adept at wedging themselvesinto lobster pots, to eat the catch and bait.Much has been made of the danger of this and other wobbegongs to people, often to the exclusion of everything else of theirlife history. This species has been known to bite people that step on it or put their feet near its mouth, and can and will bitewhen molested or provoked, as when speared or caught by line or nets. These sharks can inflict severe lacerations, andthere is one report of a fisher losing his foot to a spotted wobbegong that was disturbed in a rock pool. At least for thisspecies, inflicting fatalities on people is virtually unknown and requires verification in other wobbegongs. The strong jawsand jaw musculature, and (unlike nurse sharks) large and effective impaling teeth of these wobbegongs, coupled with theirtendency to hold on after biting, makes them a minor hazard to unwary explorers of tide-pools, fishermen and divers, but thesharks otherwise appear to be relatively unaggressive and sluggish when unprovoked, as when a diver views themunderwater. However, placing one’s limb near the head of a wobbegong may be inviting trouble, as the shark may bite eitherfrom mistaking the limb as a smaller prey item, or in response to being cornered. Several unprovoked and provoked biteincidents on people (including some on divers near the bottom and well above it) by Australian wobbegongs (probablyincluding this species), and even a few cases of wobbegongs biting boats, have been reported in the literature, but it is oftendifficult to determine which species was involved or what the precise circumstances were that led to the incident.Wobbegongs of all sizes, but especially larger individuals, should be treated with due respect.This species is displayed in large public aquaria in Europe, the United States, and probably Australia and is viewed by diversin Australia.Local Names: Wobbegong, Carpet shark, Kumohada-oose or Kumohada-ôse (Japan).Remarks: Extra-Australian records for this species require confirmation according to Last and Stevens (1998) and in thewriter’s estimation.Literature: Ogilby and McCulloch (1908); Garman (1913); Whitley (1940); Fowler (1941); Matsubara (1955); Chen (1963);Garrick and Schultz (1963); Stead (1963); Marshall (1965); Grant (1972); Compagno (1984); Nakaya and Shirai (1984);Michael (1993); Last and Stevens (1994).

158 FAO Species Catalogue for Fishery Purposes No. 1Orectolobus ornatus (De Vis, 1883) Fig. 123Crossorhinus ornatus De Vis, 1883, Proc. Linn. Soc. New South Wales (1884), 8(2): 289, Mar. 28, 1883. Holotype: Typespecimens not mentioned by De Vis but the holotype is Queensland Museum, QM I.164 (dry mount) according toEschmeyer (1998, Cat. Fish.: CD-ROM), type locality, “Moreton Bay, etc.”, Queensland.Synonyms: Orectolobus devisi Ogilby, 1916: 181. Unnecessary replacement name for Crossorhinus ornatus De Vis,1883, presumed by Ogilby to be preoccupied by Bonaparte’s (1834) inclusion of Scyllium ornatum Gray, 1832(= Chiloscyllium plagiosum) in his new genus Orectolobus. Holotype: Queensland Museum, QM I.164 (dry mount)according to Eschmeyer (1998: CD-ROM). Orectolobus ornatus halei Whitley, 1940: 81, figs 70-71. Holotype: SouthAustralian Museum, Adelaide, SAMA 2 883 mm TL female (cast), not found in 1997 according to Eschmeyer (1998:CD-ROM), St. Vincent Gulf, South Australia.Other Combinations: Crossorhinus barbatus (not Gmelin, 1788).FAO Names: En - Ornate wobbegong; Fr - Requin-tapis paste; Sp - Tapicero ornamentado.DORSAL VIEWLATERAL VIEWUNDERSIDE OF HEADFig. 123 Orectolobus ornatusField Marks: Flattened benthic sharks with dermal lobes on sides of head, symphysial groove on chin; a stronglycontrasting, variegated colour pattern of conspicuous broad dark, dorsal saddles with light spots and conspicuous black,corrugated edges, interspaced with lighter areas and conspicuous dark, light-centred spots; also, mouth in front of eyes,long, basally branched nasal barbels, nasoral grooves and circumnarial grooves, two rows of enlarged fang-like teeth inupper jaw and three in lower jaw; first dorsal-fin origin over pelvic-fin bases.Diagnostic Features: Nasal barbels with a few branches. Five dermal lobes below and in front of eye on each side of head;dermal lobes behind spiracles unbranched or weakly branched and broad. No dermal tubercles or ridges on back.Interspace between dorsal fins longer than inner margin of first dorsal fin, about half first dorsal-fin base. Origin of first dorsalfin over about last third of pelvic-fin base. First dorsal-fin height about equal to base length. Colour: colour pattern veryconspicuous and highly variegated, dorsal surface of body with conspicuous broad, dark rectangular saddles with deeplycorrugated, prominent black-edged margins, dotted with light spots and not ocellate in appearance; interspaces betweensaddles light, with numerous broad light-centred dark blotches.Distribution: Western Pacific: ?Japan, Indonesia, New Guinea, and tropical and warm-temperate Australia (Queensland,New South Wales, Victoria, South Australia, south coast of Western Australia; absent from Tasmania, northern WesternAustralia and Northern Territory).

Sharks of the World, Vol. 2 159Habitat: A common inshore bottom shark of continentalwaters, found in bays, on alga-covered rocky areas andcoral reefs on the coast and around offshore islands, inlagoons, on reef flats and faces, and in reef channels.Favours clearer water than does the spottedwobbegong. Occurs frim the intertidal to at least 100 mdepth.Biology: This is a nocturnal shark, that rests on thebottom during the day in caves, under ledges on reefs,and in trenches, and prowls on its reef habitat at night. Itis observed singly and often in aggregations during theday, sometimes with several animals piled on top of oneanother. Ovoviviparous, with litter of at least 12 young.Feeds on bony fishes, sharks, rays, cephalopods, andcrustaceans.Size: Maximum about 288 cm. Size at birth about 20 cm.Normally maturing at about 175 cm but a Queenslandmale was mature at 63 cm (suggesting the possibility ofmore than one species included under this taxon).Interest to Fisheries and Human Impact: Interest tofisheries limited, fished as bycatch of commercial sharkfisheries off Western Australia and taken by dropline offNew South Wales. Flesh marketed but of littlecommercial value at present. Skin very tough andattractively patterned, and occasionally used for makinga good leather. Said to bite waders and fishers intidepools and occasionally biting divers. Michael (1993)suggests that large males of this species may be moreaggressive during the breeding season. Sometimes theymay swim off the bottom and approach a nearby diver, possibly as an agonistic response. This species is kept in publicaquaria in the United States and probably Australia. Conservation status uncertain, but of concern in places outsideAustralian territorial waters where it is found in places subjected to habitat degradation and uncontrolled fisheries.Local Names: Gulf wobbegong, Banded wobbegong, Carpet shark, Karakusa-ôse (Japan).Remarks: Whitley (1940) proposed a subspecies, O. ornatus halei, for the ornate wobbegongs from South Australia,separable from O. o. ornatus of more northeastern waters by differences in its colour pattern and in the dermal flaps of thehead. It remains to be seen at what level these apparent differences can be recognized.Extra-Australian records for this species (Masuda, Araga and Yoshino, 1975; Nakaya and Shirai, 1984, for Japan) requireconfirmation (Last and Stevens, 1994; see also discussion above).Literature: Ogilby and McCulloch (1908); Garman (1913); Whitley (1940); Stead (1963); Marshall (1965); Grant (1972);Masuda et al. (1975); Compagno (1984); Nakaya and Shirai (1984); Michael (1993); Last and Stevens (1994).Orectolobus wardi Whitley, 1939 Fig. 124Orectolobus wardi Whitley, 1939, Rec. Australian Mus., 20: 264. Holotype: Australian Museum, Sydney, AMS-IA-7784,Cape Keith, Melville Island, northern Australia.Synonyms: None.Other Combinations: None.

160 FAO Species Catalogue for Fishery Purposes No. 1FAO Names: En - Northern wobbegong; Fr - Requin-tapis savetier; Sp - Tapicero zapatilla.DORSAL VIEWLATERAL VIEWUNDERSIDE OF HEADFig. 124 Orectolobus wardiField Marks: Flattened benthic sharks with dermal lobes on sides of head, symphysial groove on chin, variegated but rathersombre colour pattern of rounded, ocellate dark dorsal saddles with entire edging and light margins, interspaced with broaddusky areas without spots or reticular lines; also, mouth in front of eyes, long, basally branched nasal barbels, nasoralgrooves and circumnarial grooves, two rows of enlarged fang-like teeth in upper jaw and three in lower jaw.Diagnostic Features: Nasal barbels without branches. Two dermal lobes below and in front of eye on each side of head;dermal lobes behind spiracles unbranched and broad. No dermal tubercles or ridges on back. Interspace between dorsalfins longer than inner margin of first dorsal fin, about half first dorsal-fin base. Origin of first dorsal fin over about last fourth ofpelvic-fin base. First dorsal-fin height about equal to base length. Colour: colour pattern variegated but dull and sombrecompared to most other wobbegongs, dorsal surface of body with small, rounded, ocellate, light-edged saddle marks withentire margins, separated from each other by broad, dusky spaces without spots or broad reticulated lines.Distribution: Western South Pacific: Confined toAustralian waters (Queensland, Northern Territory andWestern Australia).Habitat: A little-known but possibly common tropicalinshore bottom shark of the Australian northerncontinental shelf. Occurs on shallow-water reefs in waterless than 3 m deep, often in turbid areas.Biology: A nocturnal shark, inactive during the day,sometimes seen with its head under a ledge. Probablyovoviviparous. Presumably feeds on bottominvertebrates and fishes, but diet unrecorded.Size: Maximum to at least 63 cm and possibly 100 cm; a45 cm male was mature.Interest to Fisheries and Human Impact: Interest tofisheries none at present. Conservation status unknown.Local Names: Northern wobbegong, North Australian wobbegong.Literature: Whitley (1939, 1940); Marshall (1965); Compagno (1984); Michael (1993); Last and Stevens (1994).click for next page

click for previous pageSharks of the World, Vol. 2 161Orectolobus sp. A Last and Stevens, 1994 Fig. 125Orectolobus sp. A Last and Stevens, 1994, Sharks Rays Australia: 128, pl. 26.Synonyms: None.Other Combinations: None.FAO Names: En - Western wobbegong; Fr - Requin-tapis sombre; Sp - Tapicero occidental.LATERAL VIEWDORSAL VIEWFig. 125 Orectolobus sp. AUNDERSIDE OF HEADField Marks: Flattened benthic sharks with dermal lobes on sides of head, symphysial groove on chin; a stronglycontrasting, variegated colour pattern of conspicuous broad dark, dorsal saddles with light spots and deeply corrugatededges but without conspicuous black margins, interspaced with lighter areas and conspicuous light, dark-centred spots butwithout numerous light O-shaped rings; also, mouth in front of eyes, long, basally branched nasal barbels, nasoral groovesand circumnarial grooves, two rows of enlarged fang-like teeth in upper jaw and three in lower jaw; first dorsal-fin origin overrear half of pelvic-fin bases.Diagnostic Features: Nasal barbels with one small branch. Four dermal lobes below and in front of eye on each side ofhead; dermal lobes behind spiracles unbranched or weakly branched and slender. Low dermal tubercles or ridges presenton back in young, lost in adults. Interdorsal space somewhat shorter than inner margin of first dorsal fin, about one-fourth offirst dorsal-fin base. Origin of first dorsal fin over aboutlast third of pelvic-fin base. First dorsal-fin height aboutthree-fourths of base length. Colour: colour pattern veryconspicuous and highly variegated, dorsal surface ofbody with conspicuous broad, dark rectangular saddleswith deeply corrugated margins, not black-edged, dottedwith light spots but without numerous O-shaped lightrings; saddles not ocellate in appearance; interspacesbetween saddles light, with numerous broad darkblotches.Distribution: Eastern Indian Ocean, Australia(temperate west coast of Western Australia from CapeLeeuwin to Coral Bay, but possibly not extending intomore southern waters or into tropical Western Australia).

162 FAO Species Catalogue for Fishery Purposes No. 1Habitat: Inshore on the continental shelf of Western Australia, on reefs and in seagrass. Depths unreported, but probably inwater from the intertidal to less than 100 m.Biology: Poorly known. Ovoviviparous, presumably feeds on invertebrates and small fish.Size: Maximum about 200 cm. Size at birth about 22 cm; an 85 cm male was mature.Interest to Fisheries and Human Impact: Interest to fisheries minimal, taken as a bycatch in small quantities by theWestern Australian shark fishery and used for human consumption. Viewed by divers off Western Australia. Conservationstatus uncertain but should be monitored because of its limited geographic range and presumably limited bathymetric range.Remarks: See remarks above in the family account on similarities of this species to Sutorectus tentaculatus.Literature: Compagno (1984); Michael (1993); Last and Stevens (1994).Sutorectus Whitley, 1939Genus: Sutorectus Whitley, 1939, Australian Zool., 9(3): 228.Type Species: Crossorhinus tentaculatus Peters, 1864, by original designation.Number of Recognized Species: 1.Synonyms: None.Diagnostic Features: Head rather narrow, its greatest width slightly less than distance from snout tip to first gill openings.Chin smooth, without a beard of dermal lobes. Dermal lobes of sides and front of head small, short, unbranched, and formingisolated groups that are broadly separated from one another, in 4 to 6 pairs. Nasal barbels simple and unbranched. Mouthnarrow, width about 9% of total length. Dorsal surface of head, body and precaudal tail, and dorsal-fin bases, with rows oflarge, conspicuous dermal tubercles, resembling warts. Trunk moderately broad, width across pectoral-fin insertionsconsiderably less than head length. Precaudal tail rather long, distance from pelvic-fin insertion to lower caudal-fin originmuch greater than head length. Pectoral and pelvic fins small and widely spaced from each other, distance from pectoral-fininsertions to pelvic-fin origins about twice length of pectoral-fin bases and somewhat greater than pelvic-fin lengths fromorigins to free rear tips. Interspace between first and second dorsal fins much shorter than first dorsal-fin inner margin andless than a fifth of first dorsal-fin base. Dorsal fins low and long, height of first dorsal fin about half its base length, length offirst dorsal-fin base greater than pelvic-fin length from origin to free rear tip. Origin of first dorsal fin in front of midbases ofpelvic fins. Colour: dorsal surface with a colour pattern of jagged-edged broad dark saddles and scattered dark spots on alight background, no reticulating narrow lines with spots at their junctions.Remarks: Whitley (1939) proposed the genus Sutorectus on the simple nasal barbels, tuberculate back, and narrowerinterdorsal space of the type and only species, S. tentaculatus (Peters, 1864). This genus was recognized by Whitley(1940), Bigelow and Schroeder (1948), Whitley and Pollard (1980), Compagno (1984), Dingerkus (1986), and Last andStevens (1994), but was considered a synonym of Orectolobus by Stead (1963) and Applegate (1974). The writer wasinclined to support Applegate’s classification (Compagno, 1973) prior to examining specimens of Sutorectus tentaculatus,but found additional characters by which this species may be distinguished from typical Orectolobus. As noted above (seeremarks under family), Orectolobus sp. A. approaches S. tentaculatus in certain features of its morphology.Sutorectus tentaculatus (Peters, 1864) Fig. 126Crossorhinus tentaculatus Peters, 1864, Monatsb. Akad. Wiss. Berlin: 123. Syntypes: Zoologisches Museum, Museum fürNaturkunde der Humboldt-Universität, Berlin, ZMB 5073, a 750 mm female, and ZMB 5264, a 430 mm female (alcohol),according to Paepke and Schmidt (1988, Mitt. Zool. Mus. Berlin, 64(1): 163) and Eschmeyer (1998, Cat. Fish.: CD-ROM),from Adelaide, South Australia.Synonyms: None.Other Combinations: Orectolobus tentaculatus (Peters, 1864).

Sharks of the World, Vol. 2 163FAO Names: En - Cobbler wobbegong; Fr - Requin-tapis cordonnier; Sp - Tapicero zapatudo.DORSAL VIEWLATERAL VIEWField Marks: A rather slender wobbegong, less flattened than most; with a few slender dermal lobes on sides of head,simple, unbranched nasal barbels, symphysial groove on chin; conspicuous warty tubercles in rows on the dorsal surface ofthe body and dorsal fin bases; dorsal fins very low and long, with heights half their base lengths, first dorsal-fin origin in frontof pelvic-fin midbases; striking variegated colour pattern of broad dark, dorsal saddles with jagged, corrugated edges,interspaced with light areas with irregular dark spots; also, mouth in front of eyes, nasoral grooves and circumnarial groovespresent, two rows of enlarged fang-like teeth in upper jaw and three in lower jaw.Diagnostic Features: See genus Sutorectus above.Distribution: Western South Pacific: Confined toAustralian waters (west coast of temperate WesternAustralia from Houtman Abrolhos southeast to Adelaide,South Australia).Habitat: A little-known but probably common inshorebottom shark of temperate continental waters, on rockyand coral reefs, and in seaweeds. Depths not recorded.Biology: Biology almost unknown. Presumablyovoviviparous and preying on bottom invertebrates andfishes.Size: Maximum recorded 92 cm. Size at birth about22 cm; near full-term young, still with sizeable yolk sacs,were 18 cm long. Males mature at about 65 cm. Said togrow as large as the spotted wobbegong (Orectolobusmaculatus) according to Stead (1963), but possibly byconfusion with Orectolobus ornatus or some other largewobbegong.Interest to Fisheries and Human Impact: Interest to fisheries none at present. Conservation status uncertain but needs tobe monitored because of its limited range.Local Names: Cobbler carpet shark, Cobbler shark.Fig. 126 Sutorectus tentaculatusLiterature: Ogilby and McCulloch (1908); Garman (1913); Whitley (1940); Fowler (1941); Stead (1963); Compagno (1984);Last and Stevens (1994).

164 FAO Species Catalogue for Fishery Purposes No. 12.3.4 Family HEMISCYLLIIDAEFamily: Subfamily Hemiscylliinae Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 407, 408 (Family ScylliorhinoidaeGill, 1862). Also subfamily Hemiscylliinae Fowler, 1934, Proc. Acad. Nat. Sci. Philadelphia, 85: 238 (Family Orectolobidae).Type Genus: Hemiscyllium Müller and Henle, in Smith, 1837.Number of Recognized Genera: 2.Synonyms: Subfamily Chiloscylliinae Gill, 1862b: 407, 408 (Family Scylliorhinoidae Gill, 1862). Type genus: ChiloscylliumMüller and Henle, 1837. Family Cheiloscyllium Hasse, 1879: 54. Type genus: Cheiloscyllium Hasse, 1879, =Chiloscyllium Müller and Henle, 1837. Family Hemiscylliidae Whitley, 1940: 68. Independently proposed as a separatefamily. Type genus: Hemiscyllium Müller and Henle, in Smith, 1837:86.FAO Names: En - Bamboo sharks, Longtailed carpet sharks; Fr - Requins-chabot; Sp - Bamboas.Field Marks: Small, slender sharks with nasoral grooves, perinasal grooves, short barbels, small transverse mouths in frontof eyes, dorsolateral eyes, large spiracles below eyes, no lateral skin flaps on head; two spineless dorsal fins, the seconddorsal-fin origin well ahead of the anal-fin origin, a long, low, keel-like rounded anal fin separated from the lower caudal originby a narrow notch, and a long precaudal tail much greater than the head and body length.Diagnostic Features: Head narrow to moderately broad and cylindrical to somewhat flattened, without lateral flaps of skin.Snout broadly to narrowly rounded or slightly pointed. Eyes dorsolaterally situated on head and with strong subocular ridgesbelow them. Eyes without movable upper eyelids or subocular pockets. Spiracles large and subequal in size or somewhatlarger than eyes, without prominent raised external rims; spiracles somewhat below and behind eyes. Gill slits small, fifth gillslit overlapping fourth; internal gill slits without filter screens. Nostrils with short pointed barbels, circumnarial folds andcircumnarial grooves present around outer edges of incurrent apertures. Nasoral grooves long and strongly developed.Mouth small, nearly transverse, and subterminal on head. Lower lip not trilobate and without lateral orolabial groovesconnecting edge of lip with medial ends of lower labial furrows, without a longitudinal symphysial groove on chin. Lowerlabial furrows extending medially nearly to symphysis and connected medially by a mental groove or groove and flap. Teethnot strongly differentiated in upper and lower jaws, with symphysial teeth not enlarged nor fang-like. Tooth row count 26 to35/21 to 32. Teeth with a strong medial cusp, with or without a pair of short lateral cusplets, and with weak labial root lobes.Teeth orthodont with a central pulp cavity and no plug of osteodentine. Body cylindrical or slightly depressed, with or withoutridges on sides. Precaudal tail longer than body. Caudal peduncle without lateral keels or precaudal pits. Pectoral fins small,broad and rounded. Pectoral fins aplesodic and with fin radials not expanded into fin web. Pectoral propterygium large andseparate from mesopterygium and metapterygium or fused with mesopterygium; pectoral-fin radial segments three at most,and with longest distal segments 0.3 to 0.4 times the length of longest proximal segments. Pelvic fins somewhat smaller orabout as large as dorsal fins and subequal to or much larger than anal fin, nearly as large or as large as pectoral fins and withanterior margins 0.6 to about 1.1 times pectoral-fin anterior margins. Claspers without mesospurs, claws or dactyls. Dorsalfins equal-sized. First dorsal-fin origin varying from over pelvic-fin bases to behind them, insertion well behind the pelvic-finrear tips. Anal fin somewhat smaller than second dorsal fin, with broad base, broadly rounded keel-like apex, origin behindsecond dorsal-fin insertion, and insertion separated by a narrow notch much less than base length from lower caudal-finorigin. Caudal fin horizontally elongated and not crescentic, weakly heterocercal with its upper lobe hardly elevated abovethe body axis; dorsal caudal-fin margin less than a fifth as long as the entire shark. Caudal fin with a strong terminal lobe andsubterminal notch but without a ventral lobe, preventral and postventral margins not differentiated and forming a continuouscurve. Vertebral centra with well-developed radii but no annuli. Total vertebral count 151 to 192, monospondylous precaudalcount 32 to 41, diplospondylous precaudal count 57 to 90, diplospondylous caudal count 55 to 72, and precaudal count 89 to129. Cranium narrow and not greatly expanded laterally. Medial rostral cartilage moderately long and not reduced to a lownubbin. Nasal capsules elevated and not greatly depressed or fenestrated, internarial septum high and compressed. Orbitswith small foramina for preorbital canals, medial walls not fenestrated around the optic nerve foramina. Supraorbital crestspresent on cranium but not laterally expanded nor pedicellate. Suborbital shelves moderately broad and not greatlyreduced. Cranial roof unfenestrated, with isolated frontoparietal fenestrae, or with a continuous fenestra from the anteriorfontanelle to the parietal fossa. Basal plate of cranium with separate pairs of carotid and stapedial foramina. Adductormandibulae muscles of jaws with two or three divisions. Preorbitalis muscles extending onto posterodorsal surface ofcranium. No anterodorsal palpebral depressor, rostromandibular, rostronuchal or ethmonuchal muscles. Valvular intestineof ring type with 12 to 20 turns. Development oviparous, eggs laid in elliptical egg capsules. Size small with adults betweenabout 43 and 107 cm total length with most below 1 m; young may hatch at about 9 to 17 cm. Colour pattern of dark saddlesand bars and dark or light spots present, or colour plain.Distribution: Bamboo sharks are small inshore bottom sharks of continental waters of the Indo-West Pacific, ranging fromMadagascar in the west to Japan, Philippines, and the Australian region in the east. One genus (Hemiscyllium) is centredon Australia, New Guinea and the Indo-Australian Archipelago, in the western Pacific (with an apparent outlier in theSeychelles), but the second (Chiloscyllium) is wide-ranging over the entire range of the family.Habitat: Bamboo sharks commonly occur in the intertidal, in tidepools on rocky or coral reefs close inshore, sometimes inwater sufficient only to cover them, and on soft bottoms inshore and offshore in open and enclosed bays.

Sharks of the World, Vol. 2 165Biology: Bamboo sharks are common to abundant and frequently kept in captivity, but their biology is poorly known at best.They have slender trunks and tails and strong, muscular, leg-like paired fins ideal for clambering on reefs and in crevices.These sharks are small, mostly less than 1 m maximum length. Michael (1993) suggested that the large epaulette spotsseen on Hemiscyllium species may serve as eyespots to intimidate predators approaching them from above. At least someand probably all of the species are oviparous, depositing eggs on the bottom in oval egg cases. Colour patterns of the youngare often strikingly different and bolder than adults and suggest different habitat preferences or habits. Food of these sharksis little known, but includes small bottom fishes, cephalopods, shelled molluscs, and crustaceans.Interest to Fisheries and Human Impact: Hemiscyllium species are little-utilized for fisheries apart from the aquariumtrade, but Chiloscyllium species are commonly caught in small-scale artisanal and commercial fisheries and by bottomtrawlers in the western Pacific and East-Central Indian Ocean. In some countries such as Thailand large catches ofChiloscyllium are occasionally landed. Longtailed carpet sharks are ideal for aquaria, because they are small, oftencolourful and pleasingly marked, very hardy and can live over a decade in captivity and reproduce there. Several of thespecies figure in the aquarium trade and are displayed in captivity in public aquaria worldwide. Despite their importance tofisheries and the aquarium trade the conservation status of longtailed carpet sharks is poorly known, and they may bethreatened by overfishing, bad fisheries practices and habitat modification, including the alteration and destruction of coralreefs. Some of the species are rare, have limited geographic, bathymetric, and habitat distributions, and live in poorly knownand poorly-protected areas. There is an urgent need for investigations on the biology and conservation status of thesesharks, particularly those that figure heavily in fisheries and those rarities that live in threatened and restricted habitats.Remarks: The arrangement of this family follows Garman (1913), Whitley (1940, 1967), Fowler (1941), Compagno (1984),Dingerkus (1986), Last and Stevens (1994), and especially the comprehensive review and revision of the family byDingerkus and DeFino (1983).Literature: Müller and Henle (1838d); Bleeker (1852); Dumeril (1853, 1865); Günther (1870); Regan (1908a); Smith (1913);Garman (1913); Whitley (1940, 1967); Stead (1963); Fowler (1941, 1967a); Compagno (1973, 1984, 1988); Applegate(1984); Dingerkus and DeFino (1983); Dingerkus (1986); Michael (1993); Last and Stevens (1994).Key to Genera:1a. Nostrils subterminal on snout; preoralsnout long, mouth closer to eyes thansnout tip; eyes and supraorbital ridgeshardly elevated; no black hood onhead or large dark spot or spots onsides of body above pectoral fins(Fig. 127a and b) . . . . . . . Chiloscyllium1b. Nostrils terminal on snout; preoralsnout short, mouth closer to snout tipthan eyes; eyes and supraorbitalridges prominently elevated; a largedark spot or spots on sides of bodyabove pectoral fins, or a black hoodon head (Fig. 128a and b) . . Hemiscylliuma) UNDERSIDEOF HEADnostrils subterminalno black hood or dark spotsFig. 127 Chiloscylliumnostrils terminalblack hood and spotsb) LATERAL VIEWa) UNDERSIDEOF HEADb) LATERAL VIEWFig. 128 HemiscylliumChiloscyllium Müller and Henle, 1837Genus: Chiloscyllium Müller and Henle, 1837a, Ber. K. preuss. Akad. wiss. Berlin, 2: 112; Müller and Henle, 1837b, Arch.Naturg. 3: 395 (no species mentioned); Müller and Henle, in Smith, 1837, Proc. Zool. Soc. London, 5: 85 (one species,Scyllium plagiosum Bennett, 1830); Müller and Henle, 1838a, Mag. Nat. Hist., new ser., 2: 34; Müller and Henle, 1838b,L’Institut, 6: 64; Müller and Henle, 1838c, Arch. Naturg. 4: 83 (no species mentioned); Müller and Henle, 1838d, Syst.Beschr. Plagiost., pt. 1: 17 (five species, no type allocation).Type Species: Scyllium plagiosum Bennett, 1830, by subsequent monotypy of Müller and Henle, in Smith, 1837, Proc.Zool. Soc. London, pt.5: 85; also by subsequent designation of Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 408, as“Chiloscyllium plagiosum Mül. and Henle”.

166 FAO Species Catalogue for Fishery Purposes No. 1Number of Recognized Species: 7.Synonyms: Genus Chyloscyllium Dumeril, 1853: 125. Apparent error for Chiloscyllium Müller and Henle, 1837. GenusCheiloscyllium Hasse, 1879: 51, 55 (name only); Hasse, 1882: 276. Probable emendation of Chiloscyllium Müller andHenle, 1837, as Hasse consistently used Cheiloscyllium in his work. Genus Synchismus Gill, 1862b: 407, 408. Typespecies: Chiloscyllium tuberculatum Müller and Henle, 1838, by original designation, a junior synonym of Squalus indicusGmelin, in Linnaeus and Gmelin, 1788.Diagnostic Features: Snout relatively long, preoral length over 3% of total length. Eyes and supraorbital ridges hardlyelevated. Nostrils subterminal on snout and well separated from snout tip. Nasal barbels moderately elongated, length over1.3% of total length. Mouth slightly closer to eyes than snout tip. Lower labial folds usually connected across chin by adermal fold. Pregill length more than 13.3% of total length. Preanal tail from vent to anal-fin origin usually less than 38% oftotal length. Pectoral and pelvic fins relatively thin, not heavily muscular. Pectoral-fin skeleton with propterygium separatefrom mesopterygium. Total vertebral count usually between 135 and 180. Colour pattern without a black hood on head orlarge dark spot or spots on sides of body above pectoral fins.Local Names: Bamboo sharks, Lip sharks.Remarks: The arrangement of this genus follows Dingerkus and DeFino (1983) and Compagno (1984). Nomina dubiapossibly referable to Chiloscyllium include: Squalus (Scyliorhinus) lambarda Blainville, 1816: 121 (nomen nudum).Fowler (1941: 90), suggested that this species was a possible synonym of Chiloscyllium indicum Gmelin, in Linnaeus andGmelin, 1788. Squalus (Scyliorhinus) russellianus Blainville, 1816: 121 (nomen nudum). Assumed by Fowler (1941: 86),to be based on the “Bokee Sorrah” of Russell (1803: 10, pl. 16), and a possible synonym of Chiloscyllium punctatum Müllerand Henle, 1838. Squalus (Scyliorhinus) unicolor Blainville, 1816: 121 (nomen nudum). Fowler (1941: 88) suggested thatthis species was a possible synonym of Chiloscyllium griseum Müller and Henle, 1838. Squalus (Scyliorhinus) variegatusBlainville, 1816: 121 (nomen nudum). Fowler (1941: 90) suggested that this species was a possible synonym ofChiloscyllium indicum Gmelin, in Linnaeus and Gmelin, 1788.Key to Species:1a. Body and tail very slender; anal-fin origin far behindfree rear tip of second dorsal fin, anal-finlength from origin to free rear tip subequal tolength of hypural caudal lobe from lower caudalorigin to subterminal notch; colour pattern withnumerous small dark spots, bars, and saddles ona light background (Fig. 129). . . . . Chiloscyllium indicum1b. Body and tail moderately slender to relativelystout; anal-fin origin below or close behind freerear tip of second dorsal fin, length of anal finconsiderably shorter than hypural caudal lobe;colour pattern varying from plain to barred andspotted but without numerous small dark spotsand bars (Fig. 130). . . . . . . . . . . . . . . . . . . . 2light spotsFig. 129 Chiloscyllium indicumFig. 130 Chiloscyllium plagiosum2a. Lateral ridges present on trunk; background colourof the dorsal surface of body dark with numerouslight spots. (Fig. 130) . . . Chiloscyllium plagiosum2b. Lateral ridges absent from trunk; backgroundcolour of the dorsal surface of body light, uniformor with scattered dark spots or dusky bands . . . . . . 3Fig. 131 Chiloscyllium arabicummargins concave3a. Second dorsal fin usually with a longer base thanthe first dorsal fin (Fig. 131); juveniles uniform incoloration, except for spotted fins . . Chiloscyllum arabicum3b. First dorsal fin base usually longer than the seconddorsal fin base (Fig. 132); juveniles, whereknown, with bars and spots on body . . . . . . . . . . . 4Fig. 132 Chiloscyllium punctatum

Sharks of the World, Vol. 2 1674a. Posterior margins of first and second dorsal finsdistinctly concave, free rear tips projecting; originof first dorsal fin far forward, over anterior half ofpelvic-fin bases (Fig. 132) . . . . . Chiloscyllium punctatum4b. Posterior margins of first and second dorsal finsdistinctly straight or convex, free rear tips notprojecting; origin of first dorsal fin over or behindmidlengths of pelvic-fin bases (Fig. 133) . . . . . . . . . 5margins convexFig. 133 Chiloscyllium burmensis5a. Eyes small, less than 1.25% of total length; distancefrom vent to tail tip more than 67% of totallength; prepectoral length less than 15.8% of totallength; snout to vent length less than 31% oftotal length (Fig. 133) . . . . . . . . Chiloscyllium burmensis5b. Eyes large, more than 1.25% of total length; distancefrom vent to tail tip less than 67% of totallength; prepectoral length more than 15.8% of totallength; snout to vent length more than 31% oftotal length (Fig. 134) . . . . . . . . . . . . . . . . . . . 6JUVENILEFig. 134 Chiloscyllium griseum6a. Interdorsal space usually more than 9.3% of totallength; first dorsal fin height more than 6.6% oftotal length; second dorsal fin height usuallymore than 5.8% of total length; saddle-markingsof juveniles without black edging (Fig. 134) . .. . . . . . . . . . . . . . . . . . . . Chiloscyllium griseumblack-edged saddle-markings6b. Interdorsal space usually less than 9.3% of totallength; first dorsal fin height less than 6.6% of totallength; second dorsal fin height usually lessthan 5.8% of total length; saddle-markings of juvenileswith black edging (Fig. 135). . Chiloscyllium hasseltiJUVENILEFig. 135 Chiloscyllium hasseltiChiloscyllium arabicum Gubanov, 1980 Fig. 136Chiloscyllium arabicum Gubanov, in Gubanov and Schleib, 1980, Sharks Arabian Gulf: 14, figs 6-7, pl. Type material?Persian Gulf.Synonyms: Chiloscyllium confusum Dingerkus and DeFino, 1983: 9, figs 2, 4-7, 51, 57, 61-64. Holotype: AmericanMuseum of Natural History, AMNH-44126, 408 mm TL subadult male, 18.5 km (10 mi) offshore of Calicut (Kozhikode),Kerala, India.Other Combinations: None.FAO Names: En - Arabian carpet shark; Fr - Requin-chabot camot; Sp - Bamboa arábiga.JUVENILEFig. 136 Chiloscyllium arabicum

168 FAO Species Catalogue for Fishery Purposes No. 1Field Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, greatly elongated thick precaudal tail, longand low anal fin just anterior to caudal fin, prominent predorsal and interdorsal ridges on back, dorsal fins with nearly straightposterior margins, first dorsal-fin origin opposite or just behind pelvic-fin insertions, second dorsal fin usually with a longerbase than first; no colour pattern in young and adults.Diagnostic Features: Prepectoral length 16.1 to 19.6% of total length. Snout fairly thick and rounded anteriorly. Eyesmoderately large, lengths 1.4 to 1.8% of total length. Body and tail moderately slender. Lateral ridges absent from trunk butinterdorsal and predorsal ridges prominent. Interdorsal space fairly long, nearly or over twice first dorsal-fin base and 8.7 to14.5% of total length. Snout to vent length 33.1 to 36.3% of total length; distance from vent to tail tip 61.0 to 67.6% of totallength. Dorsal fins large and subangular, subequal to or larger than pelvic fins, and without concave posterior margins andprojecting free rear tips. First dorsal-fin origin over or behind rear halves of pelvic-fin bases, first dorsal-fin base usuallyslightly shorter than second dorsal-fin base (rarely about equal), first dorsal-fin height 4.3 to 8.4% of total length. Seconddorsal-fin height 4.2 to 7.1% of total length. Origin of anal fin somewhat behind free rear tip of second dorsal fin, anal-finlength from origin to free rear tip somewhat less than hypural caudal lobe from lower caudal origin to subterminal notch,anal-fin base less than six times anal-fin height. Total vertebral count between 141 and 175 (mean = 158.2, n = 23). Intestinalvalve count unknown. Colour: colour pattern absent in adults, juveniles with light spots on fins but otherwise unmarked.Distribution: Western Indian Ocean: India, Pakistan,and the Persian Gulf between Iran and the ArabianPeninsula (including Saudi Arabia and Kuwait).Habitat: Found on coral reefs, lagoons, rocky shores,and mangrove estuaries, depths from 3 to 100 m.Biology: A common inshore to offshore bottom shark inthe Persian Gulf, especially during the summer. Biologysketchily known. Feeds on squid (Loliginidae), shelledmolluscs (Gastropoda?), crustaceans, and snake eels(Ophichthidae). It has reproduced in aquaria, and themale grabs the pectoral fin of the female while mating.Lays up to four egg-cases on coral reefs, with hatchingafter 70 to 80 days.Size: Maximum total length about 70 cm. Freeliving at 101 mm, size at hatching uncertain; matures between 45 and 54 cmlong; an adult male 58 cm.Interest to Fisheries and Human Impact: Interest to fisheries minimal at present, apparently little utilized in the PersianGulf (Gubanov and Schleib, 1980) but probably is used in Pakistan and India. The conservation status of this species isuncertain; an immediate question is whether the Gulf War had an adverse effect on populations of this shark and other Gulfspecies.Local Names: Arabian bamboo shark, Confusing bamboo shark.Remarks: This species was first described by Gubanov and Schleib (1980) but Dingerkus and DeFino (1983) described itas a separate species, Chiloscyllium confusum, without mention of C. arabicum. Compagno (1984) provisionallyrecognized C. arabicum but noted that it was apparently very close to C. punctatum. Dingerkus and DeFino’s accountclearly establishes this species as being separable from C. punctatum.Literature: Kuronuma and Abe (1972); Gubanov and Schleib (1980); Dingerkus and DeFino (1983); Compagno (1984, andunpub. data); Michael (1993); Dibelius (1993).Chiloscyllium burmensis Dingerkus and DeFino, 1983 Fig. 137Chiloscyllium burmensis Dingerkus and DeFino, 1983, Bull. American Mus. Nat. Hist. 176(1): 9, figs 3, 7, 59-60. Holotype:US National Museum of Natural History, USNM-202672, 575 (569) mm TL adult male, 15° 04’ N, 95° 51’ E, off Rangoon,Burma, 29 to 33 m. Status of holotype and additional data from Howe and Springer (1993, Smiths. Contr. Zool. [540]: 5).Synonyms: None.Other Combinations: None.

Sharks of the World, Vol. 2 169FAO Names: En - Burmese bamboo shark; Fr - Requin-chabot birman; Sp - Bamboa birmana.Field Marks: Mouth well in front of eyes, eyes very small; spineless dorsal fins far posterior on tail, greatly elongated thickprecaudal tail, long and low anal fin just anterior to caudal fin, no lateral ridges on trunk, dorsal fins with straight rear margins,first dorsal-fin origin about opposite pelvic-fin insertions; no colour pattern in adults.Diagnostic Features: Prepectoral length 15.7% of total length. Snout fairly thick and rounded anteriorly. Eyes small, length1.2% of total length. Body and tail slender. No lateral ridges on trunk, and predorsal and interdorsal ridges not prominent.Interdorsal space fairly long, nearly twice first dorsal-fin base and 11.1% of total length. Snout to vent length 30.8% of totallength; distance from vent to tail tip 67.3% of total length. Dorsal fins small and rounded-angular, subequal in size to pelvicfins, and without concave posterior margins and projecting free rear tips. First dorsal-fin origin slightly behind pelvic-fininsertions, first dorsal-fin base slightly longer than second dorsal-fin base, first dorsal-fin height 6.3% of total length. Seconddorsal-fin height 5.2% of total length. Origin of anal fin somewhat behind free rear tip of second dorsal-fin, anal-fin lengthfrom origin to free rear tip slightly less than hypural caudal lobe from lower caudal origin to subterminal notch, anal-fin basemore than six times anal-fin height. Total vertebral count 176. Intestinal valve count unknown. Colour: colour pattern absentin adult male, juvenile colour pattern unknown.Distribution: Northern Indian Ocean: Burma (Myanmar).Habitat: Unknown, collected off Rangoon (Yangon),Burma, presumably inshore and off the Irrawaddy Riverdelta.Biology: Biology virtually unknown. Eats small bonyfishes.Size: The holotype and only known specimen is a 57 cmadult male.Interest to Fisheries and Human Impact: Unknown,but presumably utilized in local fisheries. Conservationstatus unknown.Remarks: Characterization of this species fromDingerkus and DeFino (1983).Literature: Dingerkus and DeFino (1983).Fig. 137 Chiloscyllium burmensisChiloscyllium griseum Müller and Henle, 1838 Fig. 138Chiloscyllium griseum Müller and Henle, 1838d, Syst. Beschr. Plagiost., pt. 1: 19. Syntypes: “Ein Exemplar aus Malabardurch Dussumier; 6 Exemplare von Pondichery durch Belanger, in Weingeist, in Paris. Indien, Japan”. Lectotype: MuseumNational d’Histoire Naturelle, Paris, MNHN-1010, 374 mm immature male, Malabar, Kerala, India, designated by Dingerkusand DeFino, 1983, Bull. American Mus. Nat. Hist. 176(1): 12.Synonyms: None.Other Combinations: Hemiscyllium griseum (Müller and Henle, 1838).

170 FAO Species Catalogue for Fishery Purposes No. 1FAO Names: En - Grey bamboo shark; Fr - Requin-chabot gris; Sp - Bamboa gris.JUVENILEField Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, greatly elongated thick precaudal tail, longand low anal fin just anterior to caudal fin, no lateral ridges on trunk, dorsal fins with straight or convex posterior margins, firstdorsal-fin origin about opposite rear halves of pelvic-fin bases; often no colour pattern in adults, but young with transversedark bands that lack black edging.Diagnostic Features: Prepectoral length 16.5 to 19.5% of total length. Snout rounded anteriorly. Eyes moderately large,lengths 1.3 to 2.2% of total length. Body and tail fairly stout. No lateral ridges on trunk and predorsal and interdorsal ridgesnot prominent. Interdorsal space fairly short, slightly greater than first dorsal-fin base and 8.7 to 11.5% of total length. Snoutto vent length 34.5 to 38.3% of total length; distance from vent to tail tip 58.1 to 64.4% of total length. Dorsal fins fairly largeand rounded, subequal to or larger than pelvic fins, and without concave posterior margins and projecting free rear tips. Firstdorsal-fin origin over rear halves of pelvic-fin bases, first dorsal-fin base slightly longer than second dorsal-fin base, firstdorsal-fin height 6.2 to 8.2% of total length. Second dorsal-fin height 7.4 to 9.1% of total length. Origin of anal fin slightlybehind free rear tip of second dorsal fin, anal-fin length from origin to free rear tip slightly less than hypural caudal lobe fromlower caudal-fin origin to subterminal notch, anal-fin base less than six times anal-fin height. Total vertebral count from 156to 170 (mean = 161.7, n = 14). Intestinal valve count 15 to 19 (n = 2). Colour: colour pattern absent in adults but young withprominent dark saddle-marks without black edging.Distribution: Indo-West Pacific: Pakistan, India, SriLanka, Malaysia, Thailand; nominal from Indonesia,China, Japan, Philippines, and Papua New Guinea butpossibly based in part on C. hasselti.Habitat: An inshore bottom shark, on rocks and inlagoons. Depths 5 to 80 m.Biology: Common where it occurs. Oviparous, depositseggs in small oval egg cases on the bottom. Probablyfeeds mainly on invertebrates.Size: Maximum total length at least 77 cm. Freelivingindividuals down to at least 12.2 cm, size at hatchinguncertain; males maturing between 45 and 55 cm.Interest to Fisheries and Human Impact: Regularlytaken in inshore fisheries off Pakistan, India andThailand, and utilized for human food. Kept in publicaquaria in the United States. Conservation statusuncertain.Fig. 138 Chiloscyllium griseumLocal Names: Blackbanded bamboo shark; Ikan tjutjot, Ikan tjutjot pisang (Malaysia, Indonesia); Bambak gorbeh (Iran,possibly not this species); Blackbanded catshark, Catshark, Cat shark; Shimazame (Japan).Remarks: This species was described as new by Müller and Henle without mention of the earlier Scyllium griseum vanHasselt, 1823. Fowler (1941) considered van Hasselt’s species as lacking a description. Compagno (1984) listed it as atentative nomen nudum, and noted that quite likely Müller and Henle based their name Chiloscyllium griseum on the earlierScyllium griseum, as they were apparently aware of van Hasselt’s work in Java. Dingerkus and DeFino (1983) consideredvan Hasselt’s species as valid and separable from Müller and Henle’s Chiloscyllium griseum although a nomen nudum andnot available as such, and so resurrected the earliest valid name based on it, C. hasselti Bleeker, 1852 (see below).Literature: Garman (1913); Fowler (1941); Herre (1953); Gubanov and Schleib (1980); Dingerkus and DeFino (1983);Compagno (1984); Nakaya and Shirai (1984); Dibelius (1993); D. Didier (pers. comm.).click for next page

click for previous pageSharks of the World, Vol. 2 171Chiloscyllium hasselti Bleeker, 1852 Fig. 139Chiloscyllium hasselti Bleeker, 1852, Verh. Batav. Genoots. Kunst. Wet. 24: 14. Syntypes: Five specimens, 480 to 590 mm,off Java, Sumatra, and Moluccas, in British Museum (Natural History) and Rikjsmuseum van Natuurlijke Histoire, Leiden.Lectotype: British Museum (Natural History), BMNH-1867.11.28.196, 594 mm TL male, Moluccas, designated by Dingerkusand DeFino, 1983, Bull. American Mus. Nat. Hist. 176(1): 17.Synonyms: Scylia griseum van Hasselt, 1823: 315 (nomen nudum); also Scyllium griseum van Hasselt, 1824: 89, cf.Dingerkus and DeFino (1983: 17). Chiloscyllium obscurum Gray, 1851: 35 (nomen nudum). Holotype: British Museum(Natural History), BMNH-1845.6.22.122, 487 mm TL, Indonesia. Chiloscyllium indicum var. obscura Günther, 1870: 413(new combination); also Ogilby, 1888: 8 (new combination), cf. Dingerkus and DeFino (1983: 17). Chiloscyllium dolganoviKharin, 1987: 367?, fig. 5 (original in Russian), also Kharin, 1987: 67, fig. 5 (English translation). Holotype: ZoologicalInstitute, Leningrad, ZIL-46984, 9° 12’ N, 104° 34’ E, Viet Nam, 12 m.Other Combinations: None.FAO Names: En - Indonesian bamboo shark; Fr - Requin-chabot indonésien; Sp - Bamboa indonesa.JUVENILEField Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, greatly elongated thick precaudal tail, longand low anal fin just anterior to caudal fin, no lateral ridges on trunk, dorsal fins with straight or convex posterior margins, firstdorsal-fin origin about opposite rear halves of pelvic-fin bases; often no colour pattern in adults, but young with transversedark bands that have prominent black edging.Diagnostic Features: Prepectoral length 16.5 to 20.3% of total length. Snout rounded anteriorly. Eyes moderately large,lengths 1.5 to 2.4% of total length. Body and tail fairly stout. No lateral ridges on trunk and predorsal and interdorsal ridgesnot prominent. Interdorsal space fairly short, slightly greater than first dorsal-fin base and 6.6 to 11.1% of total length. Snoutto vent length 33.2 to 37.7% of total length; distance fromvent to tail tip 58.9 to 64.3% of total length. Dorsal fins fairlylarge and rounded, subequal in size to pelvic fins, andwithout concave posterior margins and projecting free reartips. First dorsal-fin origin over rear halves of pelvic-finbases, first dorsal-fin base slightly longer than seconddorsal-fin base, first dorsal-fin height 4.8 to 8.1% of totallength. Second dorsal-fin height 7.4 to 9.1% of total length.Origin of anal fin slightly behind free rear tip of seconddorsal fin, anal-fin length from origin to free rear tip lessthan hypural caudal lobe from lower caudal-fin origin tosubterminal notch, anal-fin base less than six timesanal-fin height. Total vertebral count between 156 and 175(mean = 166.1, n = 11). Intestinal valve count 15 (n = 1).Colour: colour pattern absent in adults except for duskyfins, but young with prominent saddle marks consisting ofbroad dusky patches with conspicuous black edging andseparated by light areas and blackish spots and black ordusky blotches on all fins.Distribution: Indo-West Pacific: Burma, Thailand,Malaysia, Singapore, Indonesia (Sumatra, Java, betweenCelebes and New Guinea), Borneo (Sarawak), Viet Nam.Fig. 139 Chiloscyllium hasselti

172 FAO Species Catalogue for Fishery Purposes No. 1Habitat: Probably mostly close inshore, depth to 12 m.Biology: Oviparous, eggs hatching in about December. Eggs attached to benthic marine plants.Size: Maximum total length at least 61 cm. Size at hatching between 94 and 120 mm. Males maturing between 44 and 54cm, and adult males 54 to at least 59 cm.Interest to Fisheries and Human Impact: Fished in Singapore, Thailand, and probably wherever else it occurs.Conservation status uncertain.Remarks: I follow Dingerkus and DeFino (1983) in recognizing this species, and tentatively synonymize Chiloscylliumdolganovi Kharin (1987) from Viet Nam with it. The latter species agrees with C. hasselti rather than C. griseum in its lowdorsal fins and short interdorsal space. Among other Chiloscyllium with plain or nearly plain adults, the holotype of C.dolganovi agrees with C. hasselti and differs from C. burmensis in its larger eyes, differs from C. arabicum in its shortersecond dorsal fin and low interdorsal ridges, and differs from C. punctatum in lacking concave posterior margins andproduced free rear tips on its dorsal fins.Literature: Dingerkus and DeFino (1983); Kharin (1987); Cook and Compagno (1994).Chiloscyllium indicum (Gmelin, 1788) Fig. 140Squalus indicus Gmelin, in Linnaeus and Gmelin, 1788, Syst. Nat., ed. 13, Pisces 1(3): 1503. Holotype: British Museum(Natural History), BMNH-1853.11.12.205, 274 mm TL female (dried skin), “Oceano Indico”, Gronow collection. EasternIndian Ocean. Status confirmed by Dingerkus and DeFino, 1983, Bull. American Mus. Nat. Hist. 176(1): 22.Synonyms: [?Squalus] colax Meuschen, 1781: (no pagination). Meuschen’s Index to Gronovius’ Zoophylacium wasrejected by the International Commission on Zoological Nomenclature (1950: 504) as being nonbinomial. Holotype: BritishMuseum (Natural History), BMNH-1853.11.12.205, 274 mm TL female (dried skin), “Oceano Indico”, Gronow collection.Status confirmed by Dingerkus and DeFino (1983: 22). Squalus tuberculatus Bloch and Schneider, 1801: 137. Based on“Le Squale dentele” of Lacépède, 1798: 281, pl. 11, fig. 1, no locality. Syntypes: British Museum (Natural History),BMNH-1853.11.12.205, 274 mm TL female (dried skin), “Oceano Indico”, Gronow collection (status confirmed by Dingerkusand DeFino, 1983: 22); ?BMNH-1845.7.3.143, 419 mm, Cape Sea, South Africa (locality doubtful). Also Squalus(Scyliorhinus) tuberculatus Blainville, 1816: 121 (nomen nudum), and Chiloscyllium tuberculatus Müller and Henle,1838d: 20. Squalus gronovianus Shaw, 1804: 353 (on Gronow’s Squalus caudatus; see Gronow, in Gray, 1854: 8).Holotype: British Museum (Natural History), BMNH-1853.11.12.205, 274 mm TL female (dried skin), “Oceano Indico”,Gronow collection. Status confirmed by Dingerkus and DeFino (1983: 22). ?Squalus (Scyliorhinus) dentatus Blainville,1816: 121, (nomen nudum), possibly based on “Le Squale dentele” of Lacépède, 1798: 281, pl. 11, fig. 1, no locality,according to Fowler (1941: 90). Chiloscyllium phymatodes Bleeker, 1852: 21. Holotype: Probably Rikjsmuseum vanNatuurlijke Histoire, Leiden, RMNH-7406, one of two, 410 mm female, Semarang, Java, according to Dingerkus and DeFino(1983: 22). Squalus caudatus Gronow, in Gray, 1854: 8. Holotype: British Museum (Natural History),BMNH-1853.11.12.205, 274 mm TL female (dried skin), “Oceano Indico”, Gronow collection. Status confirmed by Dingerkusand DeFino (1983: 22). Chiloscyllium colax Whitley, 1939: 228. Also Hemiscyllium colax Fowler, 1941: 89. Revival of[?Squalus] colax Meuschen, 1781.Other Combinations: Hemiscyllium indicum (Gmelin, in Linnaeus and Gmelin, 1788).FAO Names: En - Slender bamboo shark; Fr - Requin-chabot élégant; Sp - Bamboa elegante.Fig. 140 Chiloscyllium indicumUNDERSIDE OF HEAD

Sharks of the World, Vol. 2 173Field Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, greatly elongated slender precaudal tail,long and low anal fin just anterior to caudal fin, lateral ridges on trunk, dorsal fins with straight or convex posterior margins,first dorsal-fin origin opposite or just behind pelvic-fin insertions; colour pattern of numerous small dark spots, saddles, anddashes.Diagnostic Features: Prepectoral length 15.2 to 18.0% of total length. Snout narrowly rounded anteriorly. Eyes moderatelylarge, lengths 1.4 to 1.9% of total length. Body and tail very slender. A lateral ridge present on each side of trunk, and strongpredorsal and interdorsal ridges on midline of back. Interdorsal space fairly long, nearly twice first dorsal-fin base and 10 to12% of total length. Snout to vent length 32.4 to 35.1% of total length; distance from vent to tail tip 62.3 to 67.0% of totallength. Dorsal fins small and rounded, subequal to or smaller than pelvic fins, and without concave posterior margins andprojecting free rear tips. First dorsal-fin origin over or behind pelvic-fin insertions, first dorsal-fin base slightly longer thansecond dorsal-fin base, first dorsal-fin height 3.9 to 5.4% of total length. Second dorsal-fin height 3.9 to 5.1% of total length.Origin of anal fin far behind free rear tip of second dorsal fin, anal-fin length from origin to free rear tip about equal to orgreater than hypural caudal lobe from lower caudal-fin origin to subterminal notch, anal-fin base over six times anal-finheight. Total vertebral count between 166 and 170 (mean = 167.0, n = 4). Intestinal valve count 14 to 15 (n = 6). Colour:colour pattern of numerous dark brown or blackish spots, dashes and bars on light brown background present in juvenilesand adults; saddle-markings or bars of young without prominent black edging.Distribution: Indo-West Pacific: Possibly Arabian Seaand India, Sri Lanka, Singapore, Thailand, ?Malaysia,Indonesia, Viet Nam, Taiwan Island (Province of China),China, ?Republic of Korea, ?Japan, ?Philippines,?Solomon Islands.Habitat: A common but little-known inshore bottomshark. It possibly may occur in fresh water in the lowerreaches of the Perak River in peninsular Malaysia.Biology: Oviparous.Size: Maximum total length about 65 cm; freelivingindividual 13 cm but size at hatching unknown; malesmaturing between 39 and 42 cm and reaching at least54 cm; adult female 43 cm.Interest to Fisheries and Human Impact: Interest tofisheries considerable. Regularly taken in inshorefisheries in India, Sri Lanka and Thailand, and utilized forhuman food. Conservation status unknown.Local Names: Ridgebacked bamboo shark, Slender bamboo shark, Ridge back shark (Sri Lanka); Corungun sorrah,Karikkan schura (Tamil); Ettee (Malabar, India); Bokee sorrah, Ra sorrah (Telugi); Poos hee (Baluchistan); Balavala(Marathi); Nga man ing-myong (Burma); Ca cha beo, Ca nham, Ca cheo beo (Viet Nam); Yu tokele, Yu belangkas, Yubodoh, Ikan tjutjot kumbang, Ikan tjutjot tekeh (Malaysia); Chalarm gope or Frog shark, Chalarm hin or Stone shark(Thailand); Kau kang sha or Time teller shark (China); Ridgeback catshark, Catshark, Nurse shark, Tenjikuzame (Japan).Remarks: Whitley (1939) proposed that Squalus colax Meuschen, 1781 was the earliest name for this species. However,Meuschen’s work (1781) indicates that although colax apparently refers to the Gronow species later named Squalusindicus, it was never proposed in the binomial form Squalus colax but simply listed as colax. Apparently colax is notavailable for this species.Dingerkus and DeFino (1983) listed specimens from South Africa, Indonesia (Java; Semarang-Riau Archipelago; BintanIsland; Bangka Island; Muntok), Malaysia (Penang Island), Thailand (Gulf of Thailand), Singapore, and China, and did notverify the wider distribution accorded this species by previous writers. They mentioned a stuffed specimen from “Cape Seas,South Africa” collected by Dr Andrew Smith (British Museum [Natural History], BMNH-1845.7.3.143, 419 mm), but noChiloscyllium have been collected off southern Africa despite extensive collecting over the past two centuries. The specimenmight have been a waif, but it is more likely to have been procured in India or the Far East and shipped to South Africa.Literature: Garman (1913); Smith (1913); Fowler (1941); Herre (1953); Taniuchi (1979); Gubanov and Schleib (1980);Dingerkus and DeFino (1983); Compagno (1984); Nakaya and Shirai (1984).Chiloscyllium plagiosum (Bennett, 1830) Fig. 141Scyllium plagiosum Bennett, 1830, Fishes, in S. Raffles, 1830, Mem. Life Pub. Serv. Sir Thomas Stamford Raffles, London:694. Holotype: British Museum (Natural History), probably lost, Sumatra. Neotype: California Academy of Sciences,

174 FAO Species Catalogue for Fishery Purposes No. 1CAS-36046, 503 mm TL adult male, 5° 58’ S, 106° 48’ E, Java Sea, Indonesia, designated by Dingerkus and DeFino, 1983,Bull. American Mus. Nat. Hist., 176(1): 24. Also Chiloscyllium indicum var. plagiosa Günther, 1870, Cat. Fish. British Mus.,8: 412; and Chiloscyllium indicum var. plagiosum Ogilby, 1888, Cat. Fish. coll. Aust. Mus. Pt. 1, Recent PalaeichthyanFishes, White, Sydney: 8 (new combination), cf. Dingerkus and DeFino, 1983, Bull. American Mus. Nat. Hist., 176(1): 23.Synonyms: Scyllium ornatum Gray, 1830, pl. 98, fig. 2. Name and illustration only. Holotype: British Museum (NaturalHistory), BMNH-1982.2.26.1, 653 mm TL skin, China Seas. Status of holotype confirmed by Dingerkus and DeFino (1983:24). ?Scyllium plagiosum var. interruptum Bleeker, 1852: 18. Types: None known according to Eschmeyer (1998:CD-ROM), type locality Batavia (Jakarta) and Semarang (Samarang), Java, Indonesia. Chiloscyllium margaritiferumBleeker, 1863: 243. Also Chiloscyllium indicum var. margaritifera Günther, 1870: 412. Holotype (both species):Rikjsmuseum van Natuurlijke Histoire, Leiden, RMNH-7404, 250 mm immature male, Obi Island, Moluccas, Indonesia.Status of holotype confirmed by Dingerkus and DeFino (1983: 24). ?Chiloscyllium caerulopunctatum Pellegrin, 1914: 230.Holotype: Museum National d’Histoire Naturelle, Paris, MNHN-14-9, 670 mm TL female, Fort Dauphin [Taolanaro],Madagascar. Status of holotype confirmed by Bass, D’Aubrey and Kistnasamy (1975c: 49, fig. 23); and Dingerkus andDeFino (1983: 24).Other Combinations: Hemiscyllium plagiosum (Bennett, 1830).FAO Names: En - Whitespotted bamboo shark; Fr - Requin-chabot á taches blanches; Sp - Bamboa punteada.Field Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, greatly elongated thick precaudal tail, longand low anal fin just anterior to caudal fin, lateral ridges on trunk, dorsal fins with straight or convex posterior margins, firstdorsal-fin origin opposite or just behind pelvic-fin insertions; colour pattern of numerous white or bluish spots and dark spots,dark bands and saddles on a light background.Diagnostic Features: Prepectoral length 15.0 to 19.4% of total length. Snout rounded or truncated anteriorly. Eyesmoderately large, lengths 1.4 to 2.2% of total length. Body and tail fairly stout. A lateral ridge present on each side of trunk,and strong predorsal and interdorsal ridges present on midline of back. Interdorsal space short, slightly greater or less thanfirst dorsal-fin base and 9.3 to 11.6% of total length. Snout to vent length 31.1 to 35.1% of total length; distance from vent totail tip 61.8 to 67.1% of total length. Dorsal fins moderately large and rounded or angular, subequal to or larger than pelvicfins, and without concave posterior margins and projecting free rear tips. First dorsal-fin origin varying from slightly anteriorto slightly behind pelvic-fin insertions, first dorsal-fin base slightly longer than second dorsal-fin base, first dorsal-fin height5.4 to 7.3% of total length. Second dorsal-fin height 4.7 to 6.8% of total length. Origin of anal fin somewhat behind free reartip of second dorsal fin, anal-fin length from origin to free rear tip somewhat less than hypural caudal lobe from lowercaudal-fin origin to subterminal notch, anal-fin base less than six times anal-fin height. Total vertebral count between 161and 185 (mean = 173.9, n = 8). Intestinal valve count 16 to 17 (n = 2). Colour: a prominent pattern of numerous white spotson a dark brown background in juveniles and adults, with small dark spots and darker brown or blackish transverse bands orsaddles that are not conspicuously edged with black.Distribution: Indo-West Pacific: Madagascar, India, SriLanka, Malaysia (Sandakan, Sarawak, Borneo),Singapore, Thailand, Indonesia (Obi Island, Moluccas;Manado, Celebes; Sumatra), Viet Nam, China, includingTaiwan Island (Province of China), Japan, andPhilippines.Habitat: A little-known inshore bottom shark. Occurs onreefs in the tropics.Biology: Common but biology poorly known. Nocturnal,rests in reef crevices during the day, but feeds at night.Oviparous. Eats bony fishes and crustaceans.Size: Maximum total length 95 cm; possible hatchlings orfree-living specimens 9.8 to 12.5 cm; adolescent malesup to 64 cm, adult males 50 to 83 cm; an adult female95 cm.Fig. 141 Chiloscylium plagiosum

Sharks of the World, Vol. 2 175Interest to Fisheries and Human Impact: Interest to fisheries considerable. Regularly taken in inshore fisheries in India,Thailand, China, and utilized for human consumption. Marketed in Madagascar, for human consumption. A particularlyhandsome and popular aquarium species, kept in public aquaria in Europe and the United States. Conservation statusuncertain.Local Names: Whitespotted bamboo shark, Cat shark; Bluespotted bamboo shark, Requin-chabot à taches bleues,Bamboa estrellada; Ikan tjutjot matjan (Malaysia, Indonesia); Chalarm hin or Stone shark, Chalarm seour or Tiger shark,Chalarm lye or Striped shark (Thailand); Pan chu sha or Striped bamboo shark, Ta sha (China); Whitespotted cat shark,Catshark, Shiroboshi-tenjiku (Japan).Remarks: There is a question as to whether Chiloscyllium caerulopunctatum Pellegrin, 1914, known from a singlespecimen from Madagascar, is a valid species or a synonym of the wide-ranging but disjunct C. plagiosum from the northernIndian Ocean east to Japan and Philippines. Fowler (1941) synonymized C. caerulopunctatum with C. plagiosum, butBass, D’Aubrey and Kistnasamy (1975c) retained it as a valid species without comparing it with C. plagiosum or otherChiloscyllium species. After comparing the redescription of the holotype of C. caerulopunctatum in Bass, D’Aubrey andKistnasamy (1975c) with specimens of C. plagiosum Compagno (1984) tentatively recognized this species as being valid,because it seemed to differ from C. plagiosum in having larger dorsal fins, a shorter blunter snout, possibly wider mouth, anda different coloration, with smaller blue rather than white spots and lighter ground colour. Dingerkus and DeFino (1983), in acomprehensive review of the genus, synonymized C. caerulopunctatum with C. plagiosum without comment. I follow theirsynonymy here tentatively and with some reluctance, but suggest that a multivariate comparison of specimens ofMadagascar “C. caerulopunctatum” with C. plagiosum from elsewhere (including comparison of vertebral and intestinalvalve count as well as morphometrics) is required to resolve the issue. M.-L. Bauchot and G. Bianchi, pers. comm. toCompagno (1984) and Bauchot and Bianchi (1984) noted the presence of C. caerulopunctatum in fish markets inMadagascar, confirming Pellegrin’s initial record.Dingerkus and DeFino (1993) mentioned a stuffed specimen of Chiloscyllium plagiosum from “Cape Seas, South Africa”collected by Dr A. Smith (British Museum [Natural History] BMNH-1845.7.3.140, 745 mm), but extensive collecting over thepast two centuries did not reveal the presence of any Chiloscyllium in South Africa. As with Smith’s specimen of C. indicumfrom “Cape Seas” the specimen of C. plagiosum might have been a waif from the northern Indian Ocean (or Madagascar), butit is possible that the specimen was procured in India or the Far East and shipped to South Africa.Literature: Garman (1913); Pellegrin (1914); Fowler (1941); Herre (1953); Chen (1963); Bessednov (1969); Bass,D’Aubrey and Kistnasamy (1975c); Gubanov and Schleib (1980); Dingerkus and DeFino (1983); Compagno (1984); Nakayaand Shirai (1984); Bauchot and Bianchi (1984); Michael (1993); Cook and Compagno (1994); M.-L. Bauchot and G. Bianchi,(pers. comm.).Chiloscyllium punctatum Müller and Henle, 1838 Fig. 142Chiloscyllium punctatum Müller and Henle, 1838d, Syst. Beschr. Plagiost., pt. 1: 18, pl. 3, 4 (pls 4 labelled as C. griseum).Holotype: Rikjsmuseum van Natuurlijke Histoire, Leiden, probably lost, Java. Neotype: American Museum of NaturalHistory, AMNH-38153, 352 mm TL female, Batavia (Djakarta), Java, Indonesia, designated by Dingerkus and DeFino, 1983,Bull. American Mus. Nat. Hist., 176(1): 30. Fowler, 1941, Bull. U. S. Natl. Mus., (100) 13: 85 gave an earlier reference to thisspecies as “Scyllium punctatum van Hasselt, Algemein Konst., Letterbode, May 1823, p.__”, but this could not beconfirmed. Müller and Henle, 1838d, Syst. Beschr. Plagiost., pt. 1: 18, list “Scyllium punctatum. Kuhl et v. Hasselt” undertheir new Chiloscyllium punctatum.Synonyms: None.Other Combinations: Hemiscyllium punctatum (Müller and Henle, 1838).FAO Names: En - Brownbanded bamboo shark; Fr - Requin-chabot bambou; Sp - Bamboa estriada.Fig. 142 Chiloscyllium punctatumJUVENILE

176 FAO Species Catalogue for Fishery Purposes No. 1Field Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, greatly elongated thick precaudal tail, longand low anal fin just anterior to caudal fin, no lateral ridges on trunk, dorsal fins with concave posterior margins andelongated free rear tips, first dorsal-fin origin opposite anterior halves of pelvic-fin bases; usually no colour pattern in adultsbut young with bold dark transverse bands and a few dark spots.Diagnostic Features: Prepectoral length 16.2 to 18.4% of total length. Snout rounded anteriorly. Eyes moderately large,lengths 1.5 to 2.4% of total length. Body and tail moderately slender. No lateral ridges on trunk, and predorsal and interdorsalridges not prominent. Interdorsal space fairly short, slightly greater than first dorsal-fin base and 9.1 to 12.7% of total length.Snout to vent length 32.7 to 35.8% of total length; distance from vent to tail tip 61.1 to 64.4% of total length. Dorsal fins largeand angular, larger than pelvic fins, and with concave posterior margins and prominently projecting free rear tips. Firstdorsal-fin origin varying from slightly anterior to pelvic-fin origins to over anterior halves of pelvic-fin bases, first dorsal-fin baselonger than second dorsal-fin base, first dorsal-fin height 6.8 to 9.9% of total length. Second dorsal-fin height 6.4 to 8.4% oftotal length. Origin of anal fin somewhat behind free rear tip of second dorsal fin, anal-fin length from origin to free rear tip lessthan hypural caudal lobe from lower caudal-fin origin to subterminal notch, anal-fin base less than six times anal-fin height.Total vertebral count between 136 and 170 (mean = 154.7, n = 6). Intestinal valve count 20 (n = 4). Colour: light brown inadults, usually without a colour pattern, but young with broad dark transverse bars and usually a scattering of small blackishspots, bars not prominently edged with black.Distribution: Indo-West Pacific: India (east coast,Andaman Islands), Malaysia, Singapore, Thailand,Indonesia (Java, Sumatra, Sulawesi, Komodo), Viet Nam,China, Taiwan (Province of China), Japan, Philippines,south coast of New Guinea (Papua-New Guinea and IrianJaya, Indonesia), north coast of Australia (NorthernTerritory, Western Australia, Queensland).Habitat: An inshore bottom shark found on coral reefs,often in tidepools, on tidal flats, and on reef faces, butprobably also present on soft bottom offshore. Found inthe intertidal down to at least 85 m.Biology: A common shark but not often seen; young hidein crevices at the bases of coral heads, and adults undertable corals. Very tenacious of life, can survive out ofwater for a long period (half a day). Oviparous, eggsdeposited in rounded egg cases about 11 by 5 cm. Feedson bottom invertebrates and possibly small fishes, but biology little-known despite its abundance in some parts of its range.Gills sometimes infested by larval isopods (praniza-larvae of the isopod Gnathia).Size: Maximum total length about 105 cm; hatchlings about 13 to 17 cm; adult males about 68 to 76 cm; an adult female62.9 cm.Interest to Fisheries and Human Impact: Regularly taken in inshore fisheries in India, Thailand, probably Singapore,Malaysia, and Philippines, and utilized for human food. It was seen in large numbers by the writer in the fish market at SamutSakon, near Bangkok, Thailand, in 1993. In Australia it is taken in beach seines and on hook-and-line and is said to prefersquid bait; it is little utilized but regarded as good eating. It may nip divers if provoked. It also is taken in the aquarium trade anddisplayed in numerous public aquaria in Australia, Canada, Mexico, Europe, and the United States; breeds in captivity. The liveyoung are particularly handsome, with brilliant black and whitish bands on the body and fins. Conservation status uncertain.Local Names: Brownbanded bamboo shark, Gray carpet shark, Brown-banded catshark or Cat shark, Brownbandedcatshark, Spotted catshark, Brown-banded bamboo shark, Brownspotted catshark, Inuzame (Japan).Remarks: Müller and Henle (1839) listed “Scyllium punctatum Kuhl and van Hasselt” under their Chiloscylliumpunctatum, but I was unable to examine Kuhl and van Hasselt’s account of their species and so could not determine ifScyllium punctatum as proposed by Kuhl and van Hasselt was a valid species and not a nomen nudum. As per Dingerkusand DeFino (1983) and Compagno (1984), Müller and Henle’s account is retained as the first valid description of thisspecies. Fowler (1967a:103) termed this species Chiloscyllium russellianum, because the Squalus (Scyliorhinus)russellianus of Blainville (1816) was “assumed as based on Bokee sorah Russell, F. of Coromandel I 1803, 10 pl. 16.Vizagapatam, India”. However, Fowler (1941) had previously hesitated to replace the extensively-used C. punctatum withBlainville’s nomen nudum.Literature: Garman (1913); Smith (1913); Whitley (1940); Herre (1953); Stead (1963); Marshall (1964); Gubanov andSchleib (1980); Whitley and Pollard (1980); Dingerkus and DeFino (1983); Compagno (1984); Nakaya and Shirai (1984);Michael (1993); Last and Stevens (1994); Cook and Compagno (1994).

Sharks of the World, Vol. 2 177Hemiscyllium Müller and Henle, 1837Genus: Hemiscyllium Müller and Henle, in Smith, 1837, Proc. Zool. Soc. London, 5: 86 (name only, with one species,“Squalus ocellatus Bloch”, equals Squalus ocellatus Bonnaterre, 1788); Müller and Henle, 1838a, Mag. Nat. Hist., new ser.,2: 34 (definition, no species); Müller and Henle, 1838c, Arch. Naturg., 4: 83 (definition, one species, Squalus ocellatus);Müller and Henle, 1838d, Syst. Beschr. Plagiost., pt. 1: 16 (definition, one species, Hemiscyllium ocellatum).Type Species: Squalus ocellatus Bloch and Schneider, 1801, by monotypy, equals S. ocellatus Bonnaterre, 1788; also bysubsequent designation of Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 408, as “Hemiscyllium ocellatum Mül. andHenle”.Number of Recognized Species: 5 or possibly 6.Synonyms: None.Diagnostic Features: Snout relatively short, preoral length less than 3% of total length. Eyes and supraorbital ridges wellelevated. Nostrils virtually terminal on snout tip. Nasal barbels very short, length less than 1.3% of total length. Mouth slightlycloser to snout tip than to eyes. Lower labial folds not connected across chin by a dermal fold. Pregill length less than 13.3%of total length. Preanal tail from vent to anal-fin origin usually more than 38% of total length. Pectoral and pelvic fins thick andheavily muscular. Pectoral-fin skeleton with propterygium fused to mesopterygium. Total vertebral count usually more than180 centra and up to 195. Colour pattern with large black or dusky epaulette spots on shoulders above pectoral fin,sometimes partially merged with dark shoulder-saddle.Local Names: Epaulette sharks, Speckled cat sharks, Yu tadek (Malaysia).Remarks: The arrangement of this genus follows Whitley (1940, 1967), Fowler (1941), Compagno (1984) and the revisionby Dingerkus and DeFino (1983). Data published in Dingerkus and DeFino suggests that Hemiscyllium species cannot bereadily distinguished by vertebral count, morphometrics or differences in fin and body shape but are easily distinguishableby coloration. Coloration does, however, vary with growth as in Chiloscyllium, and changes in colour pattern can bedramatic. Two of the five described species are known only from adults and subadults (H. strahani and H. trispeculare),and the juvenile colour pattern remains to be described for them.Dingerkus and DeFino (1983: 54, fig. 37) reported a juvenile Hemiscyllium (USNM-123025, 148 mm female) fromAmsterdam Island, Irian Jaya, New Guinea, Indonesia, that is of uncertain identity. It is distinguished by broad darktransverse stripes on a white background, a partial dark hood covering the region of the eyes and spiracles but not the snout,and a large black saddle blotch extending from the gills to the back and flank just behind the pectoral fins. It could be anundescribed species as it does not agree with the juvenile colour patterns of H. freycineti, H. hallstromi or H. ocellatum.Alternatively, it may represent the juvenile colour pattern of H. strahani (or, much less likely, H. trispeculare). In either casethe colour change with growth would be more extreme than those of other members of the genus. Resolution of the problemawaits further collections of Hemiscyllium from New Guinea.Fig. 143 Hemiscyllium sp.There is an outlier record of a Hemiscyllium from Seychelles (Dibelius, 1993: 15, as the “Seychelles bamboo shark”, H. cf.ocellatum; and Dibelius, pers. comm.), of an adult male specimen photographed by a diver (Norbert Wu) and far west of anyother records of the genus. This has no black hood-marking and has a few scattered small to large dark spots on thepreorbital snout, no white spots, large and small scattered black spots on the head, body and tail, a large conspicuous blackepaulette spot with a light margin, smaller black spots around the epaulette-spot, numerous large spots on the pectoral andpelvic fins, and spots and possibly dark bands on the underside of the tail. It matches H. freycineti and H. trispeculare inthe key below by having spots on the preorbital snout, but does not quite agree with either of them in coloration. It is closestin coloration to H. freycineti but differs in having a more prominent epaulette spot and spots and banding on the undersideof the tail. It also resembles H. hallstromi and differs from H. ocellatum in having mostly large dark spots on its body, butdiffers from both in its spotted preorbital snout, spotted pectoral fins and spotted and banded tail. It is said to reach about 100cm and was photographed at a depth of 20 to 30 m on coral branches (Dibelius, 1993). The record probably represents anundescribed species of Hemiscyllium but its status needs to be confirmed by collection and deposition of material in amuseum collection.

178 FAO Species Catalogue for Fishery Purposes No. 1Key to Species:black hood or masklarge white spots1a. Prebranchial head and snout with an abruptblack hood or mask, black spots and bands onthe underside of the head; black epaulettespots of shoulder partially merged with blackshoulder saddle; body and fins withnumerous, conspicuous small to large whitespots (Fig. 144) . . . . . . . . . . . Hemiscyllium strahani Fig. 144 Hemiscyllium strahani1b. Head and snout light to dusky, without a blackhood, underside of head uniformly light; blackepaulette spots of shoulder strongly marked;white spots inconspicuous or absent on finsand body . . . . . . . . . . . . . . . . . . . . . . . . . 22a. Preorbital snout with numerous dark spots(Fig. 145) . . . . . . . . . . . . . . . . . . . . . . . . . 32b. Preorbital snout uniformly light, without spots(Fig. 146) . . . . . . . . . . . . . . . . . . . . . . . . . 4dark spots on snoutDETAIL OF HEADFig. 1453a. Dark spots on preorbital snout mostly lessthan half eye length, body and fins coveredwith numerous small and large dark spots thatform a reticular network of light backgroundcolour between them; black epaulette spot ofshoulder large, in the form of a conspicuouswhite-ringed ocellus with a central large blackspot and two curved black marks surroundingthe posterior half of the spot (Fig. 147) . . .. . . . . . . . . . . . . . . . . . Hemiscyllium trispeculareno spots on snoutDETAIL OF HEADwhite-ringed epaulette spotFig. 1463b. Dark spots on preorbital snout mostlysubequal to eye length, dark spots on bodyand fins mostly large and sparse, not forminga reticular network of light background colour;black epaulette spots of shoulder small tomoderately large, not in the form of aconspicuous white-ringed ocellus, darkocellar spot without curved black markssurrounding its posterior half (Fig. 148) . . .. . . . . . . . . . . . . . . . . . . Hemiscyllium freycinetiFig. 147 Hemiscyllium trispecularesmall epaulette spotFig. 148 Hemiscyllium freycineti4a. All spots on body distinctly smaller than epaulettespot; epaulette spot not surrounded by largeblack spots (Fig. 149) . . . . . . . . Hemiscyllium ocellatumsmall spots on body4b. Some body spots subequal to or larger than epaulettespot; epaulette spot surrounded by largeblack spots (Fig. 150) . . . . . . . Hemiscyllium hallstromiFig. 149 Hemiscyllium ocellatumlarge spots on bodyFig. 150 Hemiscyllium hallstromi

Sharks of the World, Vol. 2 179Hemiscyllium freycineti (Cuvier, 1824) Fig. 151Scyllium freycineti Cuvier, in Quoy and Gaimard, 1824, Zoologie, Poissons, in L. de Freycinet, Voyage aut. monde l’uranieet la physicienne. Syntypes: Museum National d’Histoire Naturelle, Paris, MNHN-A.7792, 323 mm TL male, andMNHN-B.2962, 290 mm TL male, Waigiu (Waigeo), Indonesia. MNHN-A.7792 designated as lectotype by Dingerkus andDeFino, 1983, Bull. American Mus. Nat. Hist., 176(1): 38.Synonyms: Scyllium malaisianum Lesson, 1830: 94. pl. 6. Lectotype: Museum National d’Histoire Naturelle, Paris,MNHN-7767, 685 mm TL, baïe d’Offack, Waigiou, Irian Jaya, Indonesia, type status verified by Dingerkus and DeFino(1983: 38); designated as lectotype by Eschmeyer (1998: CD-ROM). Chiloscyllium malaianum Müller and Henle, 1838d:20. Holotype: Museum National d’Histoire Naturelle, Paris, 73 cm (calculated from quoted length of 27 in, assuming 27 mmper contemporary German inch) specimen, “Meerbusen von Offack. Insel Waigiou”, possibly MNHN-7767 (see above). AlsoHemiscyllium malayanum Bleeker, 1852: 6 (variant spelling).Other Combinations: Chiloscyllium freycineti (Cuvier, 1824).FAO Names: En - Indonesian speckled carpet shark; Fr - Requin-chabot grivelé; Sp - Bamboa jaspeada.Field Marks: Mouth well in front of eyes, spineless dorsal fins far posterior on tail, extremely elongated thick precaudal tail,long and low anal fin just anterior to caudal fin, large dark spots on snout, dark wide-spaced spots on body, a moderatelylarge black spot on flanks above pectoral fins, no black hood or white spots.Diagnostic Features: Colour: prebranchial head and snout without a black hood; underside of head uniformly light andwithout dark spots in adults, but with two broad dark bands in yound; several small to large dark spots mostly subequal toeye length present on preorbital snout. Black epaulette spot of shoulder moderately large, not in the form of a conspicuouswhite-ringed ocellus, nor with two or more additional round or oblong dark spots surrounding the posterior half of theepaulette spot. White spots absent from fins and body; fins and body covered with small to large and sparse dark spots thatdo not form a reticular network of light background colour between them; pectoral and pelvic fins with conspicuous darkwebs and light margins in young, changing to scattered small and large dark spots in adults. Dark saddles on dorsal surfaceand sides of tail extending as dark crossbands onto ventral surface of the preanal tail (from pelvic-fin bases to anal-fin origin)in young, but saddles and crossbands are lost in adults which have uniform light ventral surfaces on their preanal tails.Distribution: Western South Pacific: Indonesia (IrianJaya, Waigeo), Papua New Guinea (Trobriand Islandsfrom Kuia Island, Milne Bay, and east of Oro Bay).Habitat: A little-known bottom shark, occurs on coralreefs, on sand, and in seagrass in shallow water.Biology: Common in parts of New Guinea. Biology poorlyknown. It hides in reef crevices during the day and feeds atnight.Size: To at least 72 cm total length. Smallest freelivingindividual recorded 18.7 cm; males maturing between 37and 62 cm; adult female 72.2 cm.Interest to Fisheries and Human Impact: Interest tofisheries none at present. It is not known if this shark isbeing affected by the aquarium trade. Its conservationstatus urgently needs to be assessed because part of its limited range is being impacted by expanding fisheries, includingdynamite and poison fisheries that are destroying coral reefs, and possibly localized toxic pollution from riverine miningactivities.Local Names: Freckled carpet shark.Fig. 151 Hemiscyllium freycinetiJUVENILELiterature: Garman (1913); Fowler (1941); Dingerkus and DeFino (1983); Compagno (1984); Michael (1993).

180 FAO Species Catalogue for Fishery Purposes No. 1Hemiscyllium hallstromi Whitley, 1967 Fig. 152Hemiscyllium hallstromi Whitley, 1967, Australian Zool., 14(2): 178. Syntypes: Australian Museum, Sydney,AMS-I.15717-001, 730 mm adult male, and AMS-I.15584-001, 765 mm TL adult male, vicinity of Port Moresby, Papua-NewGuinea. AMS-I.15717-001 designated as lectotype by Dingerkus and DeFino, 1983, Bull. American Mus. Nat. Hist.,176(1): 40.Synonyms: None.Other Combinations: None.FAO Names: En - Papuan epaulette shark; Fr - Requin-chabot épaulette; Sp - Bamboa hombrera.JUVENILEField Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, extremely elongated thick precaudal tail,long and low anal fin just anterior to caudal fin; no dark spots on snout, dark wide-spaced spots on body, a conspicuous largeblack ocellar spot on flanks above pectoral fins, surrounded by smaller black spots, no black hood or white spots.Diagnostic Features: Colour: prebranchial head and snout without a black hood; underside of the head uniformly light andwithout dark spots; preorbital snout without spots. Black epaulette spot of shoulder strongly marked, large, in the form of aconspicuous white-ringed ocellus and two or three large round black spots surrounding the posterior and dorsal part of theocellus. White spots absent from fins and body; body and unpaired fins with small to large dark spots, some as large asepaulette spot, dark spots loose-set and not forming a reticular network of light background colour between them; pectoraland pelvic fins with conspicuous black webs and light margins in young, fading in adults to dusky, no light or dark spots onpaired fins. Dark saddles on dorsal surface and sides of tail extending as dark crossbands onto ventral surface of preanal tailin young, but saddles and crossbands lost in adults which have uniform light ventral surfaces on their preanal tails.Distribution: Western South Pacific: Papua-NewGuinea (Port Moresby Area, Torres Strait, MurrayIsland).Habitat: A little-known inshore bottom shark, possibly oncoral reefs.Biology: Essentially unknown. A 188 mm specimen(Australian Museum, Sydney, AMS IB.7938) wasreported by Dingerkus and DeFino (1983: 40) as havingbeen “born in the Taronga Zoo Park Aquarium; received1967 (one of the syntypes was father)”, but they did notspecify if the species was ovoviviparous rather thanoviparous as with other hemiscylliids.Size: To about 77 cm total length. Smallest freelivingspecimen 18.8 cm; males maturing between 47.6 and63.9 cm; two adult males 73.0 and 76.5 cm.Fig. 152 Hemiscyllium hallstromiInterest to Fisheries and Human Impact: Interest to fisheries none at present. It is not known if this shark is being affectedby the aquarium trade. Its conservation status urgently needs to be assessed because of its rarity and restricted range. It isnot known if it is being affected by pollution or destructive fisheries practices such as dynamiting or poisoning of coral reefs.Literature: Whitley (1967); Dingerkus and DeFino (1983); Compagno (1984).

Sharks of the World, Vol. 2 181Hemiscyllium ocellatum (Bonnaterre, 1788) Fig. 153Squalus ocellatus Bonnaterre, 1788, Tabl. Encyclop. Method. Trois Reg. Nat., Ichthyol., Paris: 8. Holotype: MuseumNational d’Histoire Naturelle, Paris, MNHN-1003, 353 mm TL (immature) male, “La mer du sud”, vicinity of Cookstown,Queensland, Australia. Status of type confirmed by Dingerkus and DeFino, 1983, Bull. American Mus. Nat. Hist., 176(1): 40.Synonyms: Squalus oculatus Banks and Solander in Gray, 1827: 436. Holotype: Museum National d’Histoire Naturelle,Paris, MNHN-1003, 353 mm TL (immature) male, vicinity of Cooktown, Queensland, Australia. Status of type confirmed byDingerkus and DeFino (1983: 40).Other Combinations: Scyllium ocellatum (Bonnaterre, 1788), Hemiscyllium oculatum (Gray, 1827).FAO Names: En - Epaulette shark; Fr - Requin-chabot ocellé; Sp - Bamboa ocelada.Fig. 153 Hemiscyllium ocellatumField Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, extremely elongated thick precaudal tail,long and low anal fin just anterior to caudal fin; no spots on snout, small dark wide-spaced spots on body, a conspicuouswhite-ringed black ocellus, without surrounding black spots, on flanks above pectoral fins, no black hood or white spots.Diagnostic Features: Colour: prebranchial head and snout without a black hood; underside of the head uniformly light andwithout dark spots; preorbital snout without spots. Black epaulette spot of shoulder strongly marked, large, in the form of aconspicuous white-ringed ocellus, with scattered and inconspicuous small dark spots surrounding the posterior and dorsalpart of the epaulette spot. White spots absent from fins and body; body and unpaired fins with small to medium-sized darkspots that are much smaller than the epaulette spot, dark spots loose-set and not forming a reticular network of lightbackground colour between them; pectoral and pelvic fins with conspicuous black webs and light margins in young, fading inadults, sometimes a few small dark spots on paired fins of adults. Dark saddles on dorsal surface and sides of tail extendingas dark crossbands onto ventral surface of preanal tail in young, but saddles and crossbands lost in adults which haveuniform light ventral surfaces on their preanal tails.Distribution: Western South Pacific: New Guinea(Papua New Guinea and Irian Jaya, Indonesia), Australia(Northern Territory, Western Australia, Queensland, NewSouth Wales), Solomon Islands, possibly also Malaysiaand Sumatra.Habitat: An abundant, small, harmless tropical sharkfound on coral reefs in shallow water, often in tidepoolsand sometimes in water barely covering it. It prefersstaghorn coral stands on reef faces, coral flats, and tidepools.JUVENILEBiology: Particularly common on the Great Barrier Reef,where it can be seen crawling and clambering about andswimming on the bottom. It is more active after dark,particularly at dusk, but also coordinates its activities withtidal cycles, preferring to feed when the tide is out. It isunafraid of people and will come up to the feet of a ‘reef-fossicker’ (a person picking over a coral reef) and pick up small fooditems disturbed by the person. When disturbed it may make frantic attempts to find shelter under coral debris and increvices, but is apparently satisfied, like the ostrich myth, if its head and pectoral region is covered and its tail is exposed.When captured and firmly held by hand the epaulette shark squirms violently without being able to escape, but may nip itscaptor.Oviparous, eggs taking about 120 days to hatch. The male grabs the female by her pectoral fin while mating.

182 FAO Species Catalogue for Fishery Purposes No. 1Eats primarily polychaete worms and small decapod crabs (Xanthidae, Portunidae and Ocypodidae), but also pistol shrimps(Alpheidae), mantis shrimps (Stomatopoda), amphipods, and teleost fishes (including dragonettes, Callyonymidae); alsoshelled molluscs. Young eat more worms, small fishes and amphipods than adults, which eat more crabs and shrimps, whileadults eat larger crabs than young. While hunting for food it swims near the bottom, and may stop, and wave its snout backand forth over sandy bottom to locate prey (possibly using both electroreception and olfaction). In shallow water on sandypatches between coral heads the epaulette shark may arch its body, force its snout into the sand, and vigorously thrash itstail out of water while seeking and rooting out prey, then stops and masticates its catch.This species is frequently infested with praniza larvae of gnathiid isopods. The parasites have a preference for the cloacaand clasper regions, but are also found in the buccal and branchial cavities. Heupel and Bennett (1999) believe the parasitesdo not adversely affect the health of the host.Size: Maximum total length about 107 cm. Hatchlings are about 15 cm long; smallest freeliving specimen recorded at 16.7cm; males maturing between 59 and 62 cm, with immature or adolescents up to 62 cm and adults as small as 59 cm; an adultfemale was 64.3 cm.Interest to Fisheries and Human Impact: Interest to fisheries limited at present. Its importance to the aquarium tradeneeds to be assessed. It is a popular aquarium shark and is displayed in numerous public aquaria in Australia, Canada, andthe United States. The conservation status of this shark is uncertain, but at least part of its habitat is protected in Australia. Itis one of the sharks observed by divers and reef-fossickers on the Great Barrier Reef. It may nip people when provoked.Local Names: Epaulette shark, Itar shark.Remarks: Although this species has been reported from a fairly wide range in the Indo-Australian Archipelago, the review ofDingerkus and DeFino (1983) listed specimens only from Australia (mostly from Queensland but also northwestern Australiaand New South Wales) and Solomon Islands. Last and Stevens (1994) mapped its distribution as including tropical Australiaand both coasts of New Guinea. Nominal records from Malaysia and Sumatra (Stead, 1963) need confirmation.Literature: Garman (1913); Whitley (1940); Fowler (1941); Stead (1963); Marshall (1964); Whitley and Pollard (1980);Grant (1982); Dingerkus and DeFino (1983); Compagno (1984); Michael (1993); Last and Stevens (1994); Heupel andBennett (1998, 1999).Hemiscyllium strahani Whitley, 1967 Fig. 154Hemiscyllium strahani Whitley, 1967, Australian Zool., 14(2): 176, fig. 1. Holotype: Australian Museum, Sydney,AMS-IB.7938, 735 mm adult female, vicinity of Port Moresby, New Guinea. Status of type confirmed by Dingerkus andDeFino, 1983, Bull. American Mus. Nat. Hist., 176(1): 46.Synonyms: None.Other Combinations: None.FAO Names: En - Hooded carpet shark; Fr - Requin-chabot moine; Sp - Bamboa capuchona.Fig. 154 Hemiscyllium strahaniField Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, extremely elongated thick precaudal tail,long and low anal fin just anterior to caudal fin; white spots on body, and unique black hood on head.Diagnostic Features: Colour: juvenile coloration unknown; prebranchial head and snout of adults with a unique black‘executioner’s hood’; black spots and bands present on the underside of the head; no discrete small dark spots on snout.Black epaulette spot of shoulder partially merged with black shoulder saddle, no white ring or posterior curved spotssurrounding it. Body and fins with numerous, conspicuously small to large white spots on dark saddles and blotches; noreticular network of fine dark spots on body and fins; pectoral and pelvic fins with conspicuous black webs and white-spottedmargins and webs. Dark saddles on dorsal surface and sides of preanal tail extending onto ventral surface.

Sharks of the World, Vol. 2 183Distribution: Western South Pacific: Papua-NewGuinea (Port Moresby area and Massas (Masas)Island).Habitat: A little-known inshore bottom shark of singularand unique appearance, found on coral reefs on reeffaces and flats and favouring areas with abundant hardcorals. It may occur down to 18 m deep but Michael(1993) mostly observed it at depths of 3 to 13 m.Biology: Apparently moderately common but withbiology little-known. It is nocturnal, and hides in crevicesand under table corals during the day. The holotypesurvived seven years in the Taronga Park Zoo aquariumamong a breeding colony of Hemiscyllium ocellatum,but it apparently did not hybridize with them.Size: To about 80 cm total length. Formerly known onlyfrom two adult museum specimens (Dingerkus and DeFino, 1983), an adult male 59.4 cm long and an adult female 73.5 cm.Michael (1993) observed this species on night dives and reported specimens 75 to 80 cm long.Interest to Fisheries and Human Impact: Interest to fisheries none at present. It is not known if this very attractive littleshark is being affected by the aquarium trade. Its conservation status urgently needs to be assessed because of its rarityand because its limited range could be subject to problems including destructive reef fisheries and localized toxic pollution.Remarks: Dingerkus and DeFino (1983) reported a juvenile Hemiscyllium from Irian Jaya, Indonesia, that is of uncertainidentity but could be the young of this species (see discussion above).Literature: Whitley (1967); Dingerkus and DeFino (1983); Compagno (1984); Michael (1993).Hemiscyllium trispeculare Richardson, 1843 Fig. 155Hemiscyllium trispeculare Richardson, 1843, Icones Piscium, Pl. Rare Fish: 5, pl. 1, fig. 2. Holotype: Apparently not saved,Turtle Island, northwest Australia. Neotype: British Museum (Natural History), BMNH-1953.5.10.1, 576 mm adult male,northwestern Australia, the basis of a redescription of this species by Richardson, 1846, Zool. Erebus Terror, 2, Fishes:43-44, pl. 28, and designated as neotype by Dingerkus and DeFino, 1983, Bull. American Mus. Nat. Hist., 176(1): 51.Synonyms: None.Other Combinations: Chiloscyllium trispeculare (Richardson, 1843).FAO Names: En - Speckled carpet shark; Fr - Requin-chabot marquéterie; Sp - Bamboa moteada.Fig. 155 Hemiscyllium trispeculareField Marks: Mouth well in front of eyes; spineless dorsal fins far posterior on tail, extremely elongated thick precaudal tail,long and low anal fin just anterior to caudal fin; small dark spots on snout, numerous dark close-set spots forming a reticularpattern on body, a conspicuous white-ringed large black ocellus on flanks above pectoral fins, partly surrounded by smallerblack spots, no black hood or white spots.Diagnostic Features: Colour: juvenile coloration unknown; prebranchial head and snout of adults without a black hood;underside of the head uniformly light and without dark spots; numerous small dark spots mostly less than half eye lengthpresent on preorbital snout. Black epaulette spot of shoulder strongly marked, large, in the form of a conspicuous

184 FAO Species Catalogue for Fishery Purposes No. 1white-ringed ocellus and two curved black marks surrounding the posterior half of the spot. White spots absent from fins andbody; fins and body covered with numerous small and large, densely spaced dark spots that form a reticular network of lightbackground colour between them; pectoral fins with dusky webs and numerous small dark spots. Dark saddles on dorsalsurface and sides of preanal tail extending as dark crossbands onto ventral surface.Distribution: Western South Pacific: ?Indonesia(Moluccas), Australia (Northern Territory, WesternAustralia, Queensland).Habitat: A tropical continental shelf shark that is foundon coral reefs in shallow water, including tide pools.Biology: A common small shark, with biology poorlyknown. Often observed under table corals.Size: Maximum total length 79 cm. Size at hatchingunknown; adult males 57 to 64 cm total length; adultfemale 56 cm. Possibly a smaller species than H.ocellatum.Interest to Fisheries and Human Impact: Interest tofisheries none at present. The importance of this shark tothe aquarium trade needs to be assessed. Theconservation status of this shark is uncertain, but at leastpart of its habitat is protected in Australia.Local Names: Speckled carpet shark, Speckled catshark or Cat shark.Remarks: Indonesian (Moluccas) records of this species need confirmation.Literature: Garman (1913); Fowler (1941); Stead (1963); Whitley and Pollard (1980); Grant (1982); Dingerkus and DeFino(1983); Compagno (1984); Michael (1993); Last and Stevens (1994).2.3.5 Family STEGOSTOMATIDAEFamily: Subfamily Stegostomatinae Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 407, 408 (Family ScylliorhinoidaeGill, 1862). Also subfamily Stegostomatinae Fowler, 1934, Proc. Acad. Nat. Sci. Philadelphia, 85: 239 (FamilyOrectolobidae).Type Genus: Stegostoma Müller and Henle, 1837.Number of Recognized Genera: 1.Synonyms: Family Stegostomatidae Applegate, 1974: 748. Type genus: Stegostoma Müller and Henle, 1837.Independently proposed as a family. Family Stegastomatidae Last and Stevens, 1994: 138. Apparent error forStegostomatidae.FAO Names: En - Zebra sharks; Fr - Requins zébres; Sp - Tiburones acebrados.Diagnostic Features: Head broad, conical and somewhat flattened, without lateral flaps of skin. Snout very broadlyrounded or truncated. Eyes laterally situated on head and without strong subocular ridges below them. Eyes withoutmovable upper eyelids or subocular pockets and ridges. Spiracles large and subequal to eyes, without prominent raisedexternal rims; spiracles behind but not below eyes. Gill slits small, fifth gill slit overlapping fourth; internal gill slits without filterscreens. Nostrils with short pointed barbels but without circumnarial folds and grooves around incurrent apertures. Nasoralgrooves long and strongly developed. Mouth moderately large, nearly transverse and subterminal on head. Lower liptrilobate and with lateral orolabial grooves connecting edge of lip with medial ends of lower labial furrows, no longitudinalsymphysial groove on chin. Lower labial furrows ending medially far lateral to symphysis, not connected medially by amental groove or groove and flap. Teeth not strongly differentiated in upper and lower jaws, with symphysial teeth notenlarged nor fang-like. Tooth row count 28 to 33/22 to 32. Teeth with a strong medial cusp, a pair of short lateral cusplets, andweak labial root lobes. Teeth orthodont with a central pulp cavity and no plug of osteodentine. Body cylindrical, with strongridges on sides. Precaudal tail shorter than body. Caudal peduncle without lateral keels or precaudal pits. Pectoral fins large,broad and rounded. Pectoral fins semiplesodic and with fin radials partly expanded into fin web. Pectoral propterygium smalland separate from mesopterygium and metapterygium; pectoral-fin radial segments three to nine, and with longest distalsegments up to 1.3 times the length of longest proximal segments. Pelvic fins smaller than first dorsal fin but larger than

Sharks of the World, Vol. 2 185second dorsal fin and as large or larger than anal fin, much smaller than pectorals and with anterior margins 0.4 to 0.6 timesthe pectoral-fin anterior margins. Claspers poorly known but probably without mesospurs, claws or dactyls. Dorsal fins withsecond dorsal much smaller than first. First dorsal-fin origin expanded well ahead of pelvic-fin origins and with insertionabout over pelvic-fin bases. Anal fin larger than second dorsal fin, with broad base, angular apex, origin about oppositesecond dorsal-fin midbase or insertion, and insertion separated by a space or narrow notch much less than base length fromlower caudal-fin origin. Caudal fin greatly elongated horizontally and not crescentic, weakly heterocercal with its upper lobeat a low angle above the body axis; dorsal caudal-fin margin about half as long as the entire shark. Caudal fin with a strongterminal lobe and subterminal notch but without a ventral lobe, preventral and postventral margins not differentiated andforming a continuous curve. Vertebral centra with well-developed radii. Total vertebral count 207 to 243, monospondylousprecaudal count 43 to 49, diplospondylous precaudal count 38 to 50, diplospondylous caudal count 120 to 154, andprecaudal count 81 to 101. Cranium broad and expanded laterally. Medial rostral cartilage moderately long and not reducedto a low nubbin. Nasal capsules elevated and not greatly depressed or fenestrated, internarial septum moderately high andslightly compressed. Orbits with small foramina for preorbital canals, medial walls not fenestrated around the optic nerveforamina. Supraorbital crests present on cranium and laterally expanded and pedicellate. Suborbital shelves moderatelybroad and not greatly reduced. Cranial roof solid, without a continuous fenestra from the anterior fontanelle to the parietalfossa. Basal plate of cranium with a pair of stapedial foramina widely separated from medial carotid foramina. Adductormandibulae muscles of jaws with two divisions. Preorbitalis muscles extending onto posterodorsal surface of cranium. Noanterodorsal palpebral depressor, rostromandibular, rostronuchal or ethmonuchal muscles. Valvular intestine of ring typewith 18 turns. Development oviparous. Size large with adults between 147 and 233 cm and exceptionally over 300 cm totallength while young are hatched at about 20 to 36 cm. A unique colour pattern of black saddles with light lines in young,changing to dark spots on a light background in juveniles and adults.Remarks: The arrangement of this family follows Compagno (1973, 1984) and Applegate (1974), in recognizing amonotypic family Stegostomatidae for the highly distinctive zebra shark, Stegostoma fasciatum (Herman, 1783). Dingerkus(1986) included Stegostoma in an aggregate family Rhincodontidae for ‘higher’ orectoloboids (also including Rhincodon,Pseudoginglymostoma, Ginglymostoma and Nebrius) but the earlier arrangement is followed here pending further work onthe interrelationship of orectoloboids. Dingerkus (1986) suggested that Stegostoma was the immediate sister group of thewhale shark, Rhincodon, while Compagno (1988) suggested that Stegostoma was either the immediate sister group ofPseudoginglymostoma or of a group comprising Rhincodon plus Ginglymostoma and Nebrius.The present account follows most previous researchers in recognizing a single species of zebra shark. At least one earlyauthor, Gmelin (1788), apparently recognized separate species on the strikingly distinct juvenile (Squalus tigrinus) andsubadult-adult (S. longicaudatus) colour patterns respectively. However, the zebra shark needs to be subjected to detailedmorphological and biochemical comparisons over its vast range, to determine if it is a single species.The earliest name for the zebra shark is Squalus varius Seba, 1758, which has been used by various writers (Garman,1913; Klausewitz, 1960; and Bass, D’Aubrey and Kistnasamy, 1975a) as Stegostoma varium. However, an examination ofSeba’s (1758) descriptions of fishes showed that his nomenclature was haphazardly uninomial, binomial, and polynomial,although the name of the zebra shark could be construed as binomial: “Squalus varius; naribus ori proximis; foraminibuspare oculos; spiraculis utrinque quaternis; cauda longifilis.” (Seba, 1758).In addition to the above name and diagnosis, Seba includes a long, accurate Latin description of the zebra shark, and alegend caption in French for a good illustration of a juvenile zebra shark with barred colour pattern (Seba, 1758, pl. 34, no. 1).I did not use Seba’s name previously (Compagno, 1984) or here for this shark because his nomenclature was notconsistently binomial, and the first valid usage of the name “varius” is Stegostoma varium Garman, 1913. The InternationalCommission on Zoological Nomenclature would have to make a special ruling to make the name available. Extensive usageof Stegostoma fasciatum, S. tigrinum, and other names for the zebra shark in the literature make such an actionunnecessary in the writer’s estimation.Stegostoma Müller and Henle, 1837Genus: Stegostoma Müller and Henle, 1837a, Ber. K. preuss. Akad. wiss. Berlin, 2:112.Type Species: Squalus fasciatus Bloch and Schneider, 1801, by original designation, equals Squalus fasciatus Hermann,1783.Number of Recognized Species: 1.Synonyms: Genus Stegostonea Regan, 1929: 293. Probable error for Stegostoma Müller and Henle, 1837. GenusStegastoma Herre, 1934: 10. Probable error for Stegostoma Müller and Henle, 1837. Genus Stegastoma Last and Stevens,1994: 138. Apparent error for Stegostoma Müller and Henle, 1837.Diagnostic Features: See family Stegostomatidae above.click for next page

click for previous page186 FAO Species Catalogue for Fishery Purposes No. 1Stegostoma fasciatum (Hermann, 1783) Fig. 156Squalus fasciatus Hermann, 1783, Tab. Affin.: 302. Based on Squalus varius Seba, 1758. A senior homonym of Squalusfasciatus Bonnaterre, 1788 = Poroderma africanum (Gmelin, 1788). No types according to Eschmeyer (1998, Cat. Fish.:CD-ROM). Also, Squalus fasciatus Bloch, 1785, Naturg. Ausl. Fische, 1: 19, pl. 113. Holotype: Zoologisches Museum,Museum für Naturkunde der Humboldt-Universität, Berlin, ZMB-4449, 355 mm total length male, Indian Ocean fromTranquebar, according to Paepke and Schmidt (1988, Mitt. Zool. Mus. Berlin, 64(1): 163).Synonyms: Squalus varius Seba, 1758: 105, pl. 34, fig. 1. No type locality or specimens. Name not available becauseSeba’s usage of nomenclature was not consistently binomial (see remarks above). Squalus tigrinus Pennant, 1769: 24;(nomen nudum?). Squalus tygrinus Bonnaterre, 1788: 8, pl. 8, fig. 23. Type material uncertain. Type locality: “La mer desIndes”. Squalus tigrinus Gmelin, in Linnaeus and Gmelin, 1788: 1493. Type material uncertain. Type locality: “Oceanoindico”. Squalus varius Seba, 1758 included in synonymy. This species was probably based on juveniles with a stripedcolour pattern. Squalus longicaudatus Gmelin, in Linnaeus and Gmelin, 1788: 1496. Type material uncertain, no localitygiven. Squalus varius Seba, 1758 included in synonymy. This was not strongly distinguished from S. tigrinus, but may havebeen based on post-juveniles with a spotted colour pattern. Scyllia quinquecornuatum van Hasselt, 1823: 15. Reference toSqualus varius Seba, 1758, and presumably a replacement name for it. No types listed in Eschmeyer (1998: CD-ROM).Scyllium heptagonum Rüppell, 1837: 61, pl. 17, fig. 1. Lectotype: Senckenberg Museum, Frankfurt, SMF-3152, 105 cmstuffed specimen, possibly female, Djedda, Red Sea, according to Klausewitz (1960: 290). Stegostoma carinatum Blyth,1847: 725, pl. 25, fig. 1. Type locality: India. Whereabouts of holotype unknown according to Eschmeyer (1998: CD-ROM).Squalus pantherinus Kuhl and van Hasselt, in Bleeker, 1852: 23. Name only, in synonymy of Stegostoma fasciatum Müllerand Henle, 1838. Not Scyllium pantherinum Smith, 1837 = Poroderma pantherinum. Squalus cirrosus Gronow, in Gray,1854: 46. No locality. Reference to Squalus varius Seba, 1758, and presumably a replacement name for it. No typesaccording to Eschmeyer (1998: CD-ROM). Stegostoma varium Garman, 1913: 59. Syntypes: At least two specimens, 330mm (13 in) and about 1.53 m (5 ft) mentioned without further detail. According to Eschmeyer (1998: CD-ROM) syntypesinclude Museum of Comparative Zoology, Harvard, MCZ 55-S (possibly lost), MCZ-33437, and MCZ uncat. (shrunken skinand skull). Revival of Squalus varius Seba, 1758 and first valid use of the species name. Stegostoma tigrinum naucumWhitley, 1939: 229, fig. 2. Holotype: Australian Museum, Sydney, AMS-I.4174, Hawkesbury River, New South Wales,according to Paxton et al. (1989: 92). Scyllium quinquecarinatum Fowler, 1941: 102. Error or emendation for Scylliaquinquecornuatum van Hasselt, 1823.Other Combinations: Stegostoma tigrinus or S. tigrinum (Pennant, 1769 or Gmelin, 1788), Stegostoma tygrinum(Bonnaterre, 1788).FAO Names: En - Zebra shark; Fr - Requin zèbre; Sp - Tiburón acebrado.TOOTHJUVENILEFig. 156 Stegostoma fasciatumUNDERSIDE OF HEADField Marks: Unique large sharks that combine a broad, low caudal fin about as long as the rest of the shark with nasoralgrooves, barbels, a small transverse mouth in front of the lateral eyes, two spineless dorsal fins and an anal fin, the firstdorsal fin much larger than the second and with its origin far forwards on back, prominent ridges on the sides of the body butno strong lateral keels on the caudal peduncle. Colour: colour pattern banded or spotted. Young sharks are dark brownabove, yellowish below, with vertical yellow stripes and spots breaking the dorsal coloration into dark saddles; in specimensbetween 50 and 90 cm length the saddles break up into small brown spots on a yellow background, these becoming lesslinear and more uniformly distributed with further increase in size. There is considerable variation in the colour patternbetween individuals of like size. An albino specimen was once collected.Diagnostic Features: See family Stegostomatidae above.

Sharks of the World, Vol. 2 187Distribution: Indo-West Pacific: From the east coast of South Africa (Eastern Cape and KwaZulu-Natal Provinces),Mozambique, and Madagascar north to Tanzania and east to the Red Sea, Gulf of Aden, the Maldives, the Persian Gulf,Pakistan, India, Sri Lanka, Bangladesh, Malaysia (including Sarawak, Borneo), Singapore, Indonesia (Java, MacassarStrait, Sulawesi, Dobo and Aru Islands), Thailand, Viet Nam, Kampuchea, Philippines, China, Taiwan (Province of China),Japan, New Guinea, northern Australia (Western Australia, Northern Territory, Queensland, New South Wales), NewCaledonia, and Palau.Habitat: This is an inshore tropical shark of the continental and insular shelves of the Indo-West Pacific, that is very commonon coral reefs but also occurs offshore on soft grounds. It ranges from the intertidal down to 62 m. It has been reported fromfresh water in Philippines, but this needs to be confirmed. Adults and large spotted juveniles prefer lagoons, channels andfaces of coral reefs, reef detritus and sandy places as rest areas, but the striped young are rarely seen and may prefer waterbelow 50 m.Biology: The biology of the zebra shark is sketchily known despite being relatively common and readily observed alive bydivers on coral reefs and as catches in Indo-West Pacific fish markets. The behaviour and social organization of this shark islittle known, but it has been photographed resting on sandy areas within reefs, sometimes propped up on its pectoral finsand facing a current with open mouth. According to Michael (1993), it is usually solitary, but is rarely seen in aggregations. Itapparently is rather sluggish, at least during the daytime, and is more active at night as are nurse sharks(Ginglymostomatidae) or when motivated by the presence of food. Because of its rather slender, flexible body and caudal finit is able to squirm into narrow cracks, crevices and channels while searching for food. In captivity, it spends most of its timeresting on the bottom (at least during the day), but becomes active when food is introduced into its tank.An immature male zebra shark about 1.3 m long was observed by the writer on two separate occasions in a large tank at theWaikiki Aquarium (February 2000). It sat on the bottom of its tank in the evening on one day but became highly active duringfeeding time in the early afternoon on a subsequent day. It swam about as fast as the 1.1 to 1.2 m long blacktip reef sharks(Carcharhinus melanopterus) that it was quartered with (speed estimated at 1.0 to 1.5 m per second), and during ahalf-hour’s observation stayed near the top of the tank and swam continuously. It swam strongly, with prominent anguilliformundulations of its body and tail, and showed much manoeuvring and considerable agility while swimming. The shark brokethe surface with its caudal fin on a few occasions, churning the water, but it was not obvious if it was using its tail in anyspecial way. The caudal fin was held at a low but noticeable angle to the body axis. The elongated caudal fin seems lesslikely to be used as a weapon to herd and stun small fishes than the caudal fins of threshers (Alopiidae), but could be usedduring social activities, including courtship, as well as for facilitating entry into tight spaces.Oviparous, laying eggs in large (17 cm long, 8 cm wide and about 5 cm thick), dark brown or purplish black cases with finelateral tufts of hair-like fibres, which serve to anchor the cases to the substrate. Probably lays more than one or two eggs atonce, as four fully formed, encased eggs were found in one oviduct of an adult female.Feeds primarily on molluscs (gastropods and bivalves) but also crustaceans (crabs and shrimp), small bony fishes, andpossibly sea snakes.Size: Maximum possibly 354 cm, but most adults apparently below 2.5 m. Young hatching at a size between 20 and 36 cm;males maturing between 147 and 183 cm; females maturing between 169 and 171 cm and reaching at least 233 cm.Interest to Fisheries and Human Impact: Regularly taken in inshore fisheries in Pakistan, India, Thailand, Malaysia,Taiwan (Province of China), and elsewhere where it occurs. It is caught in bottom trawls, in floating and fixed bottom gill nets,and with longlines and other line gear. It rarely takes baited hooks. The meat is utilized fresh and dried-salted for humanconsumption; livers processed for vitamins; fins dried and processed for the oriental sharkfin trade; and offal utilized forfishmeal.

188 FAO Species Catalogue for Fishery Purposes No. 1This is a hardy shark, readily kept in captivity and is an attractive and lively aquarium exhibit. It is currently kept in severalpublic aquaria in Australia, Japan, Portugal, Singapore, Spain, and the United States.The zebra shark is unaggressive when approached underwater and has not bitten people although it is sometimes harassedby divers trying to ride it. It is popular for viewing by ecotouristic divers in the Red Sea, off the Maldives, off Phuket Island,Thailand, on the Great Barrier Reef in Queensland, Australia, and elsewhere in the Indo-West Pacific. According to Michael(1993), divers hand-feed these sharks off Phuket and the sharks allow divers to physically contact them and tolerate beingstroked on the abdomen. It is uncertain if the sharks enjoy extended contact or are undergoing tonic immobility.The conservation status of the zebra shark needs assessment, as it may have declined in areas such as the Gulf of Thailandwhere it was formerly more common. Also, it may have been adversely affected by the widespread use of explosives andpoisons to fish out reefs in the eastern Indian Ocean and western Pacific as with other reef sharks. It is not known how thisshark figures in and is influenced by the international aquarium trade. Although the adults and subadults can only thrive inlarge public aquaria, as with nurse and tawny sharks, the very attractive newly-hatched young are sufficiently small to live inthe tanks of private collectors.Local Names: Zebra shark (South Africa, Sri Lanka); Tiger shark, Sea tiger, Nurse shark, Shark with tiger-like spots,Leopard shark (Australia, South Africa); Tubarào zebra (Mozambique); Variegated shark (Maldives); Baglul (Arabia);Monkey-mouth, Monkey-mouthed shark, Pollee makum, Komrasi, Oorookoolti sorrah, Potrava, Corungun sorrah (Tamil;India); Yu checkak, Yu tokek (Malay Peninsula); Ikan tjutjot matjan (Malay Samarang); Yu tokay (Malay Pinang); Chilarmseour (Tiger shark), Seaur talay (Sea tiger, Thailand); Torafuzame (Japan); Kongarasi (Telugu); Shinvala (Marathi);Butanding (Philippines, Bikol).Literature: Seba (1758); Müller and Henle (1838d); Dumeril (1865); Günther (1870); Regan (1908a); Garman (1913);Whitley (1934, 1939, 1940); Barnard (1937); Fowler (1941, 1967a); Misra (1947); Herre (1953, 1958); Lindberg and Legeza(1959); Klausewitz (1960); Teng (1962); Chen (1963); Stead (1963); Gohar and Mazhar (1964); Marshall (1965); Bessednov(1969); Shiino (1972, 1976); Nakaya (1973); Bass, D’Aubrey and Kistnasamy (1975c); Faulkner (1975); Masuda, Araga andYoshino, (1975); Fourmanoir and Laboute (1976); Grant (1982); Uchida (1982); Compagno (1984); Nakaya and Shirai(1984); Dingerkus (1986); Anderson and Ahmed (1993); Dibelius (1993); Michael (1993); Seret (1994); Last and Stevens(1994).2.3.6 Family GINGLYMOSTOMATIDAEFamily: Ginglymostomatoidae Gill, 1862b, Ann. Lyceum Nat. Hist. New York, 7(32): 393, 395, 397, 406. Emended to familyGinglymostomatidae by Gill, 1872, Smithsonian Misc. Coll. (247): 24.Type Genus: Ginglymostoma Müller and Henle, 1837.Number of Recognized Genera: 3.Synonyms: Family Ginglystomidae Jordan, 1923: 98 (emended or erroneous spelling for Ginglymostomatidae). SubfamilyNebriinae Fowler, 1934: 238 (Family Orectolobidae). Type genus: Nebrius Rüppell, 1837. Family GinglymostomatidaeWhitley, 1940: 68. Independently proposed as a separate family. Type genus: Ginglymostoma Müller and Henle, 1837.FAO Names: En - Nurse sharks; Fr - Requins nourrices; Sp - Gatas nodriza.Field Marks: Small to large sharks with nasoral grooves but no circumnarial grooves and folds, short to long barbels, smalltransverse mouths in front of eyes; small spiracles behind but not below eyes, no lateral skin flaps on head; two spinelessdorsal fins and an anal fin, the second dorsal-fin origin well ahead of the anal-fin origin, and a short precaudal tail muchshorter than the head and body.Diagnostic Features: Head broad and flattened, without lateral flaps of skin. Snout broadly rounded or truncated. Eyesdorsolaterally or laterally situated on head, with or without strong subocular ridges below them. Eyes without movable uppereyelids or subocular pockets. Spiracles much smaller than eyes, without raised external rims; spiracles behind eyes but notbelow them. Gill slits small, fifth gill slit virtually overlapping fourth; internal gill slits without filter screens. Nostrils with short tomoderately long pointed barbels but without circumnarial folds and grooves. Nasoral grooves long and strongly developed.Mouth moderately large, transverse, and subterminal on head. Lower lip trilobate or not, with or without lateral orolabialgrooves connecting edge of lip with medial ends of lower labial furrows, no longitudinal symphysial groove on chin. Lower labialfurrows extending medially but ending well lateral to symphysis and not connected medially by a mental groove or groove andflap. Teeth not strongly differentiated in upper and lower jaws, with symphysial teeth not enlarged and fang-like. Tooth rowcount 24 to 38/22 to 32. Teeth with a strong medial cusp, one to seven pairs of short lateral cusplets, and weak labial root lobes.Teeth orthodont with a central pulp cavity and no plug of osteodentine (Pseudoginglymostoma), or osteodont with a core ofosteodentine in the crown (Ginglymostoma and Nebrius). Body cylindrical or moderately depressed, without ridges on sides.Precaudal tail shorter than body. Caudal peduncle without lateral keels or precaudal pits. Pectoral fins moderately large, broadand rounded to narrow and falcate. Pectoral fins aplesodic (Pseudoginglymostoma), semiplesodic (Ginglymostoma) or fully

Sharks of the World, Vol. 2 189plesodic (Nebrius), with fin radials variably expanded into fin web or not. Pectoral propterygium moderately large andseparate from mesopterygium and metapterygium; pectoral-fin radial segments 4 to 9, and with longest distal segments 0.4to 0.7 times the length of longest proximal segments. Pelvic fins somewhat larger to somewhat smaller than dorsal fins andanal fin, much smaller than pectoral fins and with anterior margins 0.4 to 0.7 times pectoral-fin anterior margins. Clasperswithout mesospurs, claws or dactyls. Dorsal fins either equal-sized (Pseudoginglymostoma) or with second dorsal finconsiderably smaller than first (Ginglymostoma, Nebrius). First dorsal-fin origin varying from slightly anterior to pelvic-finorigins to over their bases, insertion just behind the pelvic-fin rear tips. Anal fin about as large as second dorsal fin, withbroad base and angular apex, origin about opposite second dorsal-fin origin or midbase, and insertion separated by a spacemuch less than its base length from lower caudal-fin origin. Caudal fin horizontally elongated and not crescentic,heterocercal, with its upper lobe at a moderate angle above the body axis; dorsal caudal-fin margin less than two-fifths aslong as the entire shark. Caudal fin with a strong terminal lobe and subterminal notch but without a ventral lobe or with a veryshort one, preventral and postventral margins not differentiated and forming a continuous curve (Pseudoginglymostoma)orweakly differentiated in adults (Nebrius, Ginglymostoma). Vertebral centra with well-developed radii. Total vertebral count135 to 195, monospondylous precaudal count 35 to 57, diplospondylous precaudal count 41 to 53, diplospondylous caudalcount 49 to 96, and precaudal count 85 to 103. Cranium narrow to moderately broad but not greatly expanded laterally.Medial rostral cartilage moderately long and not reduced to a low nubbin. Nasal capsules elevated and not greatlydepressed or fenestrated, internarial septum low and compressed or moderately depressed. Orbits with small foramina(Pseudoginglymostoma) or enlarged fenestra (Ginglymostoma) for preorbital canals, medial walls not fenestrated aroundthe optic nerve foramina. Supraorbital crests present on cranium and laterally expanded and pedicellate. Suborbital shelvesmoderately broad and not greatly reduced. Cranial roof either solid (Pseudoginglymostoma) or with a continuous fenestrafrom the anterior fontanelle to the parietal fossa. Basal plate of cranium with a pair of stapediocarotid foramina(Pseudoginglymostoma) or separate carotid and stapedial foramina (Ginglymostoma). Adductor mandibulae muscles ofjaws with three divisions (Ginglymostoma). Preorbitalis muscles extending onto posterodorsal surface of cranium. Noanterodorsal palpebral depressor, rostromandibular, rostronuchal or ethmonuchal muscles. Valvular intestine of ring typewith 15 to 24 turns. Development ovoviviparous as far as is known (Ginglymostoma and Nebrius). Size small to large withadults between 53 and at least 304 cm, while young are born between 27 and 79 cm (Ginglymostoma and Nebrius). Colourpattern plain or with a few dark spots in young, no saddles, reticulations or white spots.Distribution: These are common, small to large, nocturnal, inshore bottom sharks with a circumglobal distribution insubtropical and tropical continental and insular waters. Ginglymostoma currently occurs in the eastern Pacific and tropicalAtlantic, Nebrius has a wide range in continental waters of the Indo-West Pacific, while Pseudoginglymostoma has arestricted distribution in the western Indian Ocean off East Africa. They do not occur off oceanic islands far from land andhave not penetrated the Central Pacific to the Hawaiian Island chain.Habitat: Nurse sharks occur in inshore waters in depths from the intertidal down to at least 70 m. They occur on coral androcky reefs, in sandy areas, in reef lagoons, mangrove keys, and at the surf zone, usually close inshore and sometimes inwater deep enough only to cover them.Biology: Members of the genera Nebrius and Ginglymostoma are social animals that occur in small groups while restingon the bottom; they often lie atop one another in a pile.Development is ovoviviparous in Ginglymostoma, with young that are nourished primarily by yolk while in the uterus; littersof 20 to 30 young have been reported. Nebrius is ovoviviparous and practices uterine cannibalism, with a form of oophagy inwhich the developing young apparent eat large, cased, unfertilized eggs; litters are smaller, up to four. The reproductivemode of Pseudoginglymostoma is uncertain; one female laid eggs with stout egg-cases in captivity (unfertilized),suggesting that it is oviparous.These sharks cruise and clamber on the bottom with their mouths and barbels close to the substrate while searching forfood; when they contact a food item with their barbels, they quickly reverse and use their short, small mouths and largemouth cavities as a bellows to suck in their prey. The presence of small, active reef fishes in the stomachs of large,seemingly clumsy nurse and tawny nurse sharks suggest that they may stalk and suddenly suck in such items, or merelysuck them in when the prey fishes are torpid and lying on the bottom at night. Food items include a variety of bottom and reeforganisms, bony fishes, crabs, shrimp, lobsters, and other crustaceans, squid, octopi, and other molluscs, corals, seaurchins and sea squirts.Interest to Fisheries and Human Impact: The larger species of nurse sharks are or were formerly common in shallowmarine waters and are often caught in local inshore fisheries. They are utilized for human consumption, for liver oil, and fortheir thick and exceptionally tough hides, which make extremely good leather. Ginglymostoma and Nebrius are very toughand hardy sharks that can survive over a decade in captivity and are often kept in large public aquaria and oceanaria, whileyoung Ginglymostoma figure importantly in the aquarium trade. Pseudoginglymostoma is seldom kept in captivity but isalso very hardy and is more suitable for smaller aquaria. Nebrius and especially Ginglymostoma should be treated withrespect, as they will occasionally bite and clamp on to a human tormentor when provoked; their vice-like jaws may need tobe pried lose from a victim. Ginglymostoma cirratum has bitten people without provocation, but more often will bite andotherwise defend itself when harassed by divers. The small prey, small mouths and small teeth of these sharks suggest thatoccasional incidents are agonistic or defensive and not predatory. Ginglymostoma and Nebrius are popular subjects ofecotouristic dive tours.Local Names: Nurse sharks, Tubarões de leite (Mozambique).

190 FAO Species Catalogue for Fishery Purposes No. 1Remarks: This family is recognized following the works of Compagno (1973, 1982, 1984, 1988) and Applegate (1974),which revived the Ginglymostomatidae and other orectoloboid family-group taxa of Gill (1862b). The genus Nebrius wassometimes considered a synonym or subgenus of Ginglymostoma (Agassiz, 1838; Dumeril, 1853, 1865; Günther, 1870;Fowler, 1941), or Ginglymostoma was synonymized with Nebrius (Gray, 1851), Nebrius is recognized as a separate genusfrom Ginglymostoma following Garman (1913), Whitley (1940), Bigelow and Schroeder (1948), Garrick and Schultz (1963),Compagno (1973, 1984, 1988), Applegate (1974), Bass, D’Aubrey and Kistnasamy (1975c), and Dingerkus (1986).Nebrius and Ginglymostoma are usually distinguished by structural tooth characters. According to Bigelow and Schroeder(1948), Ginglymostoma has teeth with the “central cusp” largest and with several series functional, while Nebrius has teethwith “all cusps equal” (cusps as large as cusplets) and with only one or two series functional. However, Nebrius materialexamined by the writer varied considerably in cusp size, but in no instance were the cusps as small as the cusplets (cuspswere smallest in young sharks, largest in adults). The number of functional tooth series was 2 or 3 in Nebrius specimens butoverlapped Ginglymostoma cirratum with 3 or 4. Hence the arrangement and definition of these genera are revised(Compagno, 1984, and unpublished work), and the two are readily distinguished otherwise by tooth arrangement, externalmorphology, and anatomical characters (see diagnostic features of the genera).Compagno (1984) provisionally placed Ginglymostoma brevicaudatum in the same genus as G. cirratum, but noted thatthe former is strongly divergent morphologically from G. cirratum, which is closer to Nebrius ferrugineus, and differs fromthe other two species in having orthodont tooth structure, much smaller cusplets, larger cusps and narrower crowns on itsteeth, smaller barbels, more posterior first dorsal-fin origin, equal-sized dorsal fins, a shorter caudal fin, as well as otherexternal and cranial differences. Compagno (1984) suggested that G. brevicaudatum was generically distinct fromGinglymostoma proper, and, following a suggestion by D. Ward (pers. comm.), might be referable to the fossil tooth genusEostegostoma Herman and Crochard, 1977.Eostegostoma was proposed by Herman and Crochard (1977: 133) for Ginglymostoma angustum Nolf and Taverne 1977,based on fossil teeth from the Eocene of Belgium. These authors considered Eostegostoma an early stegostomatid (hencethe name), but Cappetta (1987), who recognized Eostegostoma as a genus, suggested that it was closer to Brachaelurusand fell in the family Brachaeluridae instead of Stegostomatidae. Compagno (1988) suggested that Eostegostoma wasmore Stegostoma-like than G. brevicaudatum, and that the latter requires its own genus. Dingerkus (1986) erected a newgenus, Pseudoginglymostoma, for G. brevicaudatum, which was recognized by Compagno (1988, 1999) and which isadopted here. Whatever the relationships of Eostegostoma it apparently is not congeneric with Pseudoginglymostoma.Dingerkus (1986) presented a hand-fitted cladogram of the orectoloboids, and included the genera herein placed in theGinglymostomatidae in the family Rhincodontidae along with Rhincodon and Stegostoma. In Dingerkus’ analysis his genusPseudoginglymostoma was considered the sister taxon of the other members of his Rhincodontidae, which formed twopairs of sisters, Nebrius with Ginglymostoma and Rhincodon with Stegostoma. Compagno (1988) independently produceda similar hand-fitted cladogram with two variants, placing “G. brevicaudatum”(=Pseudoginglymostoma) as the sister of the‘advanced’ orectoloboids (Stegostoma the sister of the group Rhincodon plus the sister pair Ginglymostoma and Nebrius),or, less probably with Pseudoginglymostoma as the sister of Stegostoma alone, with these taxa the sister of Rhincodonplus Ginglymostoma and Nebrius.Further work on the morphology of Pseudoginglymostoma, including its orolabial structures, neurocranium and pectoral-finskeleton, confirms its distinctiveness, and suggests that Dingerkus (1986) and Compagno (1988) were correct in placing itas the sister taxon of the advanced orectoloboids. Pending a detailed study of the morphology of the genus,Pseudoginglymostoma is retained in the Ginglymostomatidae, which may make the family paraphyletic. It may benecessary in the future to place Pseudoginglymostoma in its own family as an alternative to lumping all of the advanced andhighly divergent orectoloboids in the Rhincodontidae as per Dingerkus (1986), but as a stopgap Pseudoginglymostoma isplaced in a subfamily of its own within Ginglymostomatidae, defined as follows:Pseudoginglymostomatinae: Eyes very small and 0.8 to 1.0% of total length. Nasal barbels very short, stubby, notvery tapered, less than 1% of total length, and not reaching mouth. Lower lip not trilobate and without orolabialgrooves. Tooth rows 24 to 27/22 to 27. Teeth not compressed, tooth crown feet very narrow, cusps large, cuspletsvery small and one or two on each side; teeth with moderately broad basal ledges. Teeth orthodont and with a centralpulp cavity. Pectoral fins broadly rounded and not falcate in adults; pectoral fins semiplesodic and with radialsreaching only 55% into fin web, radial segments four. First dorsal-fin origin over or slightly behind pelvic-fin midbases.Second dorsal fin about as large as first dorsal fin. Anal fin posterior margin ends in front of lower caudal-fin origin.Caudal fin short with dorsal caudal-fin margin less than 25% of total length (adults). Total vertebral count 135 to 143,monospondylous precaudal count 35 to 37, diplospondylous caudal count 49 to 54 and 36 to 39% of total count. Jawsbroadly arcuate. Adults small, less than 1 m long.Ginglymostomatinae: Eyes small but usually over 1% of total length. Nasal barbels moderately elongated, tapered,slender and over 1% of total length, reaching past mouth. Lower lip trilobate and divided by shallow orolabial groovesconnecting mouth with lower labial furrows. Tooth rows 29 to 42/26 to 34. Teeth moderately to greatly compressed,tooth crown feet broad, cusps moderately tall to very short, cusplets moderately large and three or more on eachside, basal ledges moderately broad to very broad. Teeth osteodont, with pulp cavity filled by osteodentine. Pectoralfins semifalcate or falcate in adults, fins plesodic and with radials reaching about 80% into fin web, radial segmentsseven to eleven in longest radials. First dorsal-fin origin about over or slightly behind pelvic-fin origins. Second dorsalfin distinctly smaller than first dorsal fin. Anal free rear tip and posterior margin extending behind level of lower

Sharks of the World, Vol. 2 191caudal-fin origin. Caudal fin elongate with dorsal caudal-fin margin over 25% of total length in adults. Total vertebralcount 168 to 195, monospondylous precaudal count 48 to 57, diplospondylous caudal count 73 to 96 and 43 to 50%of total count. Adults large, 230 cm or longer.Literature: Müller and Henle (1838d); Dumeril (1865); Günther (1870); Regan (1908a); Garman (1913); Fowler (1941,1967a); Whitley (1940); Bigelow and Schroeder (1948); Stead (1963); Compagno (1973, 1984, 1988); Applegate (1974);Dingerkus (1986); Michael (1993); Last and Stevens (1994).Key to Genera:1a. Nasal barbels greatly reduced, notreaching mouth; lower lip not trilobate,no shallow orolabial groovesconnecting edge of lip with lowerlabial furrows; second dorsal and analfins about as large as first dorsal fin;caudal fin short, less than 20% of totallength (Pseudoginglymostomatinae)(Fig. 157a and b) . . Pseudoginglymostoma1b. Nasal barbels elongated, reachingmouth; lower lip trilobate, with shalloworolabial grooves connecting edge oflip with lower labial furrows; seconddorsal and anal fins much smallerthan first dorsal fin; caudal fin longer,over 25% of total length (Ginglymostomatinae)(Fig. 158a and b). . . . 22a. Eyes and gill openings dorsolateral onhead; teeth not compressed on sidesof jaw, not imbricated; pectoral, dorsaland anal fins apically rounded,pectoral fins broad and not falcate orsemifalcate (Fig. 158b) . . . Ginglymostoma2b. Eyes and gill openings lateral onhead; teeth more or less compressedon sides of jaws, forming imbricatedseries; pectoral, dorsal and anal finsapically angular, pectoral fins narrowand falcate (Fig. 159). . . . . . . . . Nebriussmall barbelsa) UNDERSIDEOF HEADlong barbelsa) UNDERSIDEOF HEADfalcate pectoralsb) LATERAL VIEWlarge 2 nd dorsaland anal finsFig. 157 Pseudoginglymostoma brevicaudatumsmall 2 nd dorsaland anal finsrounded apicesb) LATERAL VIEWFig. 158 Ginglymostoma cirratumangular apicesFig. 159 Nebrius ferrugineusGinglymostoma Müller and Henle, 1837Genus: Ginglymostoma Müller and Henle, 1837a, Ber. K. preuss. Akad. wiss. Berlin, 2: 113; Müller and Henle, 1837b, Arch.Naturg., 3: 396; Müller and Henle, 1838a, Mag. Nat. Hist., new ser., 2: 35; Müller and Henle, 1838b, L’Institut, 6:64(nospecies mentioned); Müller and Henle, 1838d, Syst. Beschr. Plagiost., pt. 1: 22 (two species, G. concolor and G. cirratum,but no type allocation); Müller and Henle, 1838, in Bonaparte, 1838, Nuov. Ann. Sci. Nat., Bologna, ser. 1, 2: 212 (onespecies mentioned, “Ginglymostoma cirrosum, Mull. et Henle”).Type Species: Squalus cirratus Gmelin, 1788, by subsequent designation of Jordan and Gilbert, 1883, Bull. U.S. Natl.Mus., 16: 18, equals S. cirratus Bonnaterre, 1788. Gill (1862b, Ann. Lyceum Nat. Hist. New York, 7[32]: 406) designated“Ginglymostoma concolor” (Müller and Henle, 1838 = Nebrius concolor Rüppell, 1837), but this would makeGinglymostoma and Nebrius synonyms. Bonaparte’s restriction of Ginglymostoma to “G. cirrosum” (a synonym of to S.cirratus) and Jordan and Gilbert’s (1883) designation are followed here. See also Bigelow and Schroeder (1948, Mem.Sears Fnd. Mar. Res., (1), 1: 180), who cited Hay’s (1902, U.S. Geolog. Surv. Bull., 179: 310) subsequent type designationof S. cirratus, and Eschmeyer (1998, Cat. Fish.: CD-ROM).Number or Recognized Species: 1.

192 FAO Species Catalogue for Fishery Purposes No. 1Synonyms: Genus Ginglimostoma Agassiz, 1838: 85. Error or emendation of Ginglymostoma. Genus GynglimostomaDumeril, 1859: 261. Probable error for Ginglymostoma. Genus Gingylostoma Springer, 1938: 13. Apparent error forGinglymostoma.Diagnostic Features: Head in dorsal or ventral view broadly arcuate in young, narrower and U-shaped in adults. Snoutbluntly wedge-shaped in lateral view, short and with mouth width about 2.3 to 2.6 times preoral length. Eyes small but usuallyover 1% of total length, positioned dorsolateral on head and with strong subocular ridges below them. Eyes with ventraledges just above level of dorsal ends of gill slits. Gill openings dorsolaterally situated on head and not or hardly visible frombelow but just reaching horizontal head rim in dorsal view. Nostrils nearly terminal on snout. Nasal barbels moderatelyelongated, tapered, slender and over 1% of total length, reaching past mouth. Lower lip trilobate and divided by shalloworolabial grooves connecting mouth with lower labial furrows. Distance between lower labial furrows about 1.5 times theirlength. Tooth rows 30 to 42/28 to 34, functional tooth series at least 7 to 9/8 to 12. Teeth moderately compressed, notimbricated but in alternate overlap pattern, functional series not separated from replacement series by toothless space.Tooth crown feet broad, cusps moderately tall, cusplets moderately large and 2 to 6 on each side, basal ledges moderatelybroad. Teeth osteodont, with pulp cavity filled by osteodentine. Body semifusiform. Lateral trunk denticles broad andrhomboid in adults, with three strong parallel ridges and a very short, blunt cusp. Pectoral fins broad and rounded-angular inyoung and semifalcate in large specimens, apices rounded. Pectoral-fin origins slightly behind to slightly in front of third gillslits. Pectoral fins plesodic and with radials reaching about 80% into fin web, radial segments 7 or 8 in longest radials. Pelvicfins rounded in young, possibly subangular in adults. Dorsal fins apically rounded. First dorsal-fin origin about over, slightly infront, or slightly behind pelvic-fin origins. Second dorsal fin distinctly smaller than first dorsal fin. Anal fin distinctly smallerthan second dorsal fin, apically rounded. Anal-fin origin about opposite, slightly behind, or slightly in front of midbase ofsecond dorsal-fin, with apex about under anal-fin base and posterior margin extending behind level of lower caudal-finorigin. Caudal fin long with dorsal caudal-fin margin over 25% of total length (adults). Caudal fin narrow and shallow withdorsal caudal-fin margin 3.2 to 3.6 times caudal-fin depth; no ventral caudal-fin lobe in young but a weak one in adults;preventral caudal-fin margin much shorter than postventral margin and 43 to 67% of it; terminal lobe short and 15 to 19% ofdorsal caudal-fin margin. Total vertebral count 168 to 175, monospondylous precaudal count 48 to 50, diplospondylouscaudal count 73 to 83 and 43 to 48% of total count. Jaws broadly arcuate. Intestinal valve count 16 to 17. Developmentovoviviparous with young nourished on yolk sac only, litter size 21 to 30. Adults 230 cm long or larger, size at birth 27 to30 cm. Colour: yellowish to grey-brown above and light whitish brown below, young with small dark, light-ringed ocellarspots and obscure dorsal saddle markings, adults and subadults without spots or saddles.Remarks: Scope of this genus is modified from Compagno (1973, 1984), and Applegate (1974), and follows Dingerkus(1986) in recognizing a single living species, Ginglymostoma cirratum. Ginglymostoma brevicaudatum is placed in thegenus Pseudoginglymostoma (see below). There are numerous fossil species of Ginglymostoma (Cappetta, 1987).Ginglymostoma cirratum (Bonnaterre, 1788) Fig. 160Squalus cirratus Bonnaterre, 1788, Tabl. Encyclop. Method. Trois Reg. Nat., Ichthyol., Paris: 7. Lectotype: MuseumNational d’Histoire Naturelle, Paris, MNHN-A.7654, 458 mm immature female, “Les mers de la Amerique.” Fromunpublished addendum to Bertin (1939b, Bull. Mus. Nat. Hist. Nat., 2e ser., 12(6): 51-98) by M.L. Bauchot (pers. comm.),also Eschmeyer (1998, Cat. Fish.: CD-ROM). Separately described as Squalus cirratus Gmelin, in Linnaeus and Gmelin,1788, Syst. Nat., ed. 13, Pisces 1(3): 1492. Types?Synonyms: Squalus punctulatus Lacépède, 1800: 119, pl. 4, fig. 3. Type locality: “Guiane”. No types known according toEschmeyer (1998: CD-ROM). Squalus cirrhatus Bloch and Schneider, 1801: 128. Based on S. cirratus Gmelin, inLinnaeus and Gmelin, 1788, possibly an emended or erroneous spelling. Also Ginglymostoma cirrhatum Jordan, 1905:533. Squalus punctatus Bloch and Schneider, 1801: 134. Based on the “Gata” of Parra, 1787: 86, pl. 34, fig. 2, from Cuba.No types known according to Eschmeyer (1998: CD-ROM). Scyllium cirrhosum Griffith and Smith, in Cuvier, Griffith andSmith, 1834: 10, pl. 30. Variant spelling? Ginglymostoma cirrosum “Müller and Henle”, in Bonaparte, 1838: 213. Variantspelling. Squalus argus Bancroft, 1832: 82. Holotype possibly in British Museum (Natural History) according to Eschmeyer(1998: CD-ROM), who indicates that this species was not characterized in the original account, but was described in anaccompanying account by Bennett (1832: 86-91). Type locality: West Indies. Ginglymostoma fulvum Poey, 1858: 342; alsoPoey, 1860, pl. 19, figs. 15-16. Holotype: 839 mm female, Cuba. Paepke and Schmidt (1988: 162) list Institut für SpezielleZoologie und Zoologisches Museum, Berlin, ZMB-5508, 965 mm TL specimen, as a possible syntype. Ginglymostomacaboverdianus Capello, 1867: 167. Cape Verde. Three syntypes according to Eschmeyer (1998: CD-ROM), but status ofthese types otherwise uncertain. Ginglymostoma cirrotum Gudger, 1914: 176. Erroneous spelling?Other Combinations: Nebrius cirratum (Bonnaterre, 1788), Scyllium cirratum (Bonnaterre, 1788).

Sharks of the World, Vol. 2 193FAO Names: En - Nurse shark; Fr - Requin nourrice; Sp - Gata nodriza.UPPER AND LOWER TEETHFig. 160 Ginglymostoma cirratumUNDERSIDE OF HEADField Marks: Moderately long barbels, nasoral grooves present but no circumnarial grooves; eyes dorsolateral, mouth wellin front of eyes, spiracles minute; precaudal tail shorter than head and body; two spineless, broadly rounded, dorsal fins andan anal fin, first dorsal fin much larger than second dorsal and anal fins, caudal fin moderately long, over one-fourth of totallength; colour yellow-brown to grey-brown, young with small dark, light-ringed ocellar spots and obscure dorsal saddlemarkings, adults and subadults without spots or saddles.Diagnostic Features: See genus Ginglymostoma above.Distribution: Western Atlantic: Rhode Island tosouthern Brazil, including United States (exceptionallyRhode Island and North Carolina, South Georgia andFlorida and Gulf coast from Florida to Texas), Mexico(Gulf of Mexico and Caribbean coasts), Bermuda,Bahamas, Turks and Caicos Islands, Cuba, Puerto Rico,Haiti, the Lesser Antilles, Belize, Jamaica, Barbados,Trinidad, French Guiana, Guyana, Panama, and Brazil(south to Rio de Janeiro). Eastern Atlantic: Cape VerdeIslands, Senegal, Cameroon to Gabon, and rarely northto Gulf of Gascony, France. Eastern Pacific: Mexico(Baja California Sur, Gulf of California) south to CostaRica, Panama, Ecuador, and Peru. The knowndistribution of the nurse shark suggests at least threegeographically isolated populations (eastern Pacific,western Atlantic and eastern Atlantic), but theirdifferentiation, if any, has yet to be studied.Habitat: This is an inshore bottom shark of the continental and insular shelves in tropical and subtropical waters, oftenoccurring at depths of1morless in the intertidal, but down to at least 12 m and off Brazil between 40 and 130 m. The nurseshark is often found on rocky and coral reefs, in channels between mangrove keys and on sand flats.Biology: This is a large nocturnal and facultatively social shark that is proverbially sluggish during the daytime butstrong-swimming and active at night; it rests on sandy bottom or in caves and crevices in rocks and coral reefs in shallowwater during the day, often in schools or aggregates of three to three dozen individuals that are close to, or even piled on oneanother while resting. Swimming speed was measured at 31 to 78 cm/sec for a 250 cm captive individual (Hussain, 1991). Inaddition to swimming near the bottom or well off it, the nurse shark can clamber on the bottom using its flexible, muscularpectoral fins as limbs. Preliminary studies suggest that the nurse shark shows a strong preference for certain day-restingsites, and repeatedly homes back to the same caves and crevices after a night’s activity. Nurse sharks fitted withconventional and sonic tags show little if any local movement, but adults have a larger range than young (Carrier, 1990). Thissuggests that groups of these animals are site-localized and vulnerable to local extirpation from overexploitation. This sharkhas been historically common or abundant in some areas where it occurs, particularly in the tropical western North Atlanticand off Tropical West Africa.Courtship and copulatory behaviour has been observed in captivity (Klimley, 1980) and studied in detail in the wild (Carrier,Pratt and Martin, 1994), and is apparently rather complex. In captivity a pair or sometimes a triplet of adults engage insynchronized parallel swimming, with the male abreast or slightly behind and below the female, but with sides nearlytouching. A pair may rest on the bottom on their bellies in parallel after bouts of parallel swimming. While parallel-swimming,the male may grab one of the female’s pectoral fins with his mouth, which in turn may induce the female to pivot 90° and rollon her back on the bottom. The male then nudges the female into a position parallel to him, swims on top of the female in

194 FAO Species Catalogue for Fishery Purposes No. 1parallel, inserts a single clasper in her vent, and then rolls on his back to lie motionless besides the inverted female withclasper still inserted. Carrier, Pratt and Martin divided nurse shark mating into five stages based on field observations offree-ranging individuals in a mating area in the Dry Tortugas, Florida: Precoupling, in which a male or group of malesapproached a female that was resting or swimming, in the latter case with parallel or tandem swimming, and with malesapproaching alongside and slightly behind the female with heads close to her pectoral fins. Coupling, in which the malegrabs the female’s pectoral fin, sometimes with two males grabbing both pectoral fins and with other males circling in closeproximity. Positioning, in which the male, or two males, roll the female onto her back, and with the male rolling and aligninghis tail and pelvic fins prior to copulation. Insertion and copulation, in which the male copulates with the female, inserting hisright clasper if holding her right pectoral, and his left clasper if holding her left pectoral, and thrusts against the female whoremains quiescent. Postcopulation, in which the male removes his claspers, releases the female’s pectoral, and either thetwo depart or lie on the bottom in parallel with ventral surfaces down. Over half the copulatory bouts involved more than onemale, with a few insertions and copulations involving two or more males.Reproduction is ovoviviparous, with intrauterine development of young being sustained primarily by the large supply of yolkin their yolk-sacs. Young are common in late spring and summer in waters off Florida, when females give birth. Numbers ofintrauterine eggs or young are 20 to 30 in a litter. The gestation period is about five to six months and it reproduces everyother year. Nursery areas are in shallow turtle-grass beds and on shallow coral reefs. Nurse sharks are slow-growing, withfree-ranging tagged juveniles (average about 126 cm long) growing at about 13 cm per year. Males may be 10 to 15 yearsold at maturity and females 15 to 20 years old (Carrier, 1990; Carrier and Luer, 1990).The nurse shark feeds heavily on bottom invertebrates such as spiny lobsters, shrimps, crabs, sea urchins, squid, octopi,marine snails and bivalves, and also fish including herring (Clupeidae), sea catfish (Ariidae), mullet (Mugilidae), parrotfish(Scaridae), surgeonfish (Acanthuridae), puffers (Tetraodontidae), and stingrays (Dasyatidae). Algae is occasionally found inits stomach and may be sucked in along with prey animals. Its small mouth and large, bellows-like pharynx allow it to suck infood items at high speed. This powerful suction feeding mechanism and its nocturnal activity pattern may allow the nurseshark to take small, active prey like bony fishes that are resting at night but would be too active and manoeuvrable for thisbig, lumbering shark to capture in the daytime. When dealing with big, heavy-shelled conchs the nurse shark flips them overand extracts the snail from its shell, presumably by grabbing its body with its teeth and by suction. It will dig under coraldetritus and in sand with its head to root out prey.Young nurse sharks have been observed resting with their snouts pointed upward and their bodies supported off the bottomon their pectoral fins; this has been interpreted as possibly providing a false shelter for crabs and small fishes that the sharkthen ambushes and eats. In captivity the nurse shark, when stimulated by food in the form of cut fish, will cruise in circlesclose to the bottom searching for the food, with its barbels touching or nearly touching the bottom; when it contacts a chunkof food, it may overshoot it but then quickly backs up and rapidly sucks it in. It may even work over vertical surfaces with itsbarbels.Size: Maximum said to be 430 cm long but most adults are under 3 m long and the largest reliably reported were 280 to 304cm. Newborn young are about 27 to 30 cm. Males mature at about 210 cm and adult males reach at least 257 cm; femalesare immature at 225 to 235 cm and mature mostly at about 230 to 240 cm (though one adult female 152 cm long has beenreported) and reach over 259 cm.Interest to Fisheries and Human Impact: This shark is commonly captured in local artisanal fisheries in some areas whereit occurs, particularly the tropical western Atlantic. It was, however, rare in a localized broad-spectrum fishery out ofGuaymas, Mexico (examined by the writer in 1974). It has been prized for its extremely tough, thick, armour-like hide, whichmakes an exceptionally good leather, but is also used fresh and salted for human consumption, as well as for liver oil and fishmeal. The stratoconidia (earstones) of this shark and other species are said to be used as a diuretic by local fishermen insouthern Brazil. It is easily captured with line gear, gill nets, fixed bottom nets and bottom trawls, and spears. It can be readilycaptured on sportsfishing tackle, but is generally regarded as being too sluggish to be much of a game fish (unlike the tawnynurse shark in Australia). Divers have sometimes speared nurse sharks, which is inane ‘sport’ because of its senselessness.The sharks are often sitting ducks for spearfishing divers on the bottom, and the modest speed of these sharks even whenactive make them no great challenge to hit. However, the toughness of these sharks may make them difficult to subdueunderwater, and a diver that spears one may receive a well-deserved bite. The nurse shark was regarded as a pest byfishermen in the Lesser Antilles because it rifled fish traps for food.The nurse shark is generally regarded as harmless to people, because of its sluggishness during the day and relatively smallteeth, and is a popular subject for ecotouristic divers, with dive sites to view this shark mostly in the Bahamas but also offBelize, Turks and Caicos, and Florida. In the Caribbean and off Florida people frequently come in contact with it underwater,and it mostly will not respond defensively when approached and usually swims away when disturbed. There have been asmall number of nonfatal, unprovoked incidents of nurse sharks biting swimmers and divers. Nonfeeding aggression,possibly related to courtship, defensive threat, or a defensive response to physical contact is possible as the motivation forsuch encounters. The small prey taken by these sharks and lack of mammalian meat in their diet suggests that predation onpeople is most improbable. In one unusual and somewhat amusing (but not to the diver) unprovoked incident a large nurseshark grabbed a diver’s chest with its teeth, then appeared to hold onto his body with its pectoral fins; the encountersuggests displaced courtship, but unfortunately the sex of the shark was not recorded. More commonly people attempt toride, spear, grab or otherwise harass this shark, or accidentally step on one while wading, and get bitten as a result.Juveniles are thought to be more prone to turn and bite than adults, which often swim away. Although its teeth are small, thejaws and associated muscles of the nurse shark are powerful and vice-like; in some instances nurse sharks have bitten

Sharks of the World, Vol. 2 195people and held on, and had to be pried loose with a tool. Hence one should treat these sharks with due respect and enjoytheir presence underwater without harassing them.Nurse sharks are very hardy and capable of surviving a wide range of temperatures and dissolved oxygen levels in captivity.They grow to adult size when obtained as young and kept in aquaria of sufficient size, and will even give birth in captivity.Specimens have been kept for 24 to 25 years. The young make interesting pets and can be trained to feed at the surface.These sharks have often been used for experimental behavioural and physiological research in captivity, for which they areexcellent subjects because of their hardiness and ability to learn. These sharks are ideal for public display in aquaria and areimportant for the aquarium trade. They are currently kept in numerous aquaria in Brazil, Europe, Mexico, the United States,and probably elsewhere.The conservation status of the nurse shark needs to be assessed throughout its range, but particularly off West Africa, partsof the tropical western Atlantic and in the eastern Pacific where intensive inshore fisheries are being pursued and thedistribution and abundance of these sharks is sketchily known. They are regarded as particularly vulnerable tooverexploitation because of their slow growth, low reproductive rate, inshore habitat, apparent site-specificity, theirpresence in areas with intense inshore fisheries, and very little catch data available. The USA was the only country reportingnurse shark catches (214 t in 1995) to FAO over the vast range of this shark, and this was only reported during the lastdecade. Apart from their vulnerability to inshore fisheries, these sharks have been increasingly captured for private andpublic aquaria, and may have declined in some areas as a result of exploitation. Carrier (1996) and Carrier and Pratt (1998)suggested that public entry should be banned during the late spring and summer at one nurse shark breeding site in the DryTortugas National Park, Florida Keys, to avoid disturbing the mating of the sharks in shallow water. These sharks are a majorasset to ecotourism in the Caribbean, and probably generate far more revenue there as live sharks viewed by divers than asfisheries products.Local Names: Nurse shark, Sand shark, Gata (Mexico, West Indies), Gata atlantica, Cacao lixa (Brazil).Literature: Gray (1851); Dumeril (1853, 1865); Günther (1870); Garman (1913); Fowler (1936, 1967a); Beebe and Tee-Van(1941a, b); Bigelow and Schroeder (1948); Cadenat (1950, 1957); Orces (1952); Springer (1960, 1963, 1973); Randall(1961, 1963, 1967, 1968, 1996); Garrick and Schultz (1963); Limbaugh (1963); Clark (1963); Clark and von Schmidt (1965);Kato, Springer and Wagner (1967); Sadowsky (1967); Böhlke and Chaplin (1968); Klimley (1974, 1980); Applegate et al.(1979); Cadenat and Blache (1981); Castro (1983); Compagno (1984, 1988); Neto, Bezerra, and Gadig (1989); Springer(1990); Carrier (1990); Carrier and Luer (1990); Michael (1993); Carrier, Pratt and Martin (1994); Bonfil (1997); Moore(1997); Carrier and Pratt (1998); Hoese and Moore (1998); McEachran and Fechhelm (1998); Castro, Woodley and Brudek(1999).Nebrius Rüppell, 1837Genus: Nebrius Rüppell, 1837, Neue Wirbel. Fauna Abyssinien gehör., Fische Rothen Meeres, (1835-1838): 62.Type Species: Nebrius concolor Rüppell, 1837, by monotypy.Number of Recognized Species: 1.Synonyms: Genus Nebrodes Garman, 1913: 56. Replacement name for Nebrius Rüppell, 1837, thought by Garman to bepreviously occupied by Nebria Latrielle, 1802 (Insecta) and Nebris Cuvier and Valenciennes, 1830 (Osteichthyes), andhence taking the same type species, Nebrius concolor Rüppell, 1837. Unjustified replacement, as Nebria and Nebris arenot senior homonyms of Nebrius.Diagnostic Features: Head in dorsal and ventral view narrow and more or less U-shaped at all stages. Snoutwedge-shaped in lateral view, moderately elongated, with preoral length about 63% of mouth width. Eyes small but usuallyover 1% of total length, situated laterally on head and with no subocular ridges below them but with strong supraocularridges above them; eyes with ventral edges below level of dorsal ends of gill slits. Gill openings laterally situated onhorizontal head rim and prominently visible from above and below. Nostrils distinctly ventral on snout. Nasal barbelsmoderately elongated, tapered, slender, over 1% of total length, and reaching past mouth. Lower lip trilobate and divided byshallow orolabial grooves connecting mouth with lower labial furrows. Distance between lower labial furrows about 2.1 timestheir length. Tooth rows 29 to 33/26 to 28, functional series 2 to 4/2 to 4. Teeth greatly compressed, with imbricate overlappattern, and functional series separated from replacement series by narrow toothless space. Tooth crown feet broad, cuspssmall, three or more moderately large cusplets on each side, and greatly widened basal ledges. Teeth osteodont and withpulp cavity filled by osteodentine. Lateral trunk denticles more or less rounded-rhomboid in adults, with four or fiveincomplete weak radial ridges and a very short, blunt cusp. Pectoral fins narrow and falcate at all stages, apically angular.Pectoral-fin origins about opposite or slightly anterior to fourth gill slits. Pectoral fins plesodic and with radials reaching about80% into fin web, radial segments 10 or 11 in longest radials. Pelvic fins angular. Dorsal fins apically angular. First dorsal-finorigin about over or slightly ahead of pelvic-fin origins. Second dorsal fin distinctly smaller than first dorsal fin. Anal fin aslarge or slightly larger than second dorsal fin, apically angular. Anal-fin origin varying from slightly in front of origin of second

196 FAO Species Catalogue for Fishery Purposes No. 1dorsal fin to about opposite its midbase, apex behind rear tip of second dorsal fin, posterior margin extends behind level oflower caudal-fin origin. Caudal fin elongate with dorsal caudal-fin margin over 25% of total length (adults); caudal fin narrowand shallow with depth 28 to 31% of dorsal caudal-fin margin; a short and weak ventral caudal-fin lobe usually present at allstages; preventral margin much shorter than postventral margin and 36 to 45% of it; terminal lobe very short and 10 to 15%of dorsal caudal-fin margin. Total vertebral count 189 to 195, monospondylous precaudal count 55 to 57, diplospondylouscount 92 to 96 and 47 to 50% of total count. Jaws narrowly arcuate. Intestinal valve count 23 to 24. Developmentovoviviparous but with yolk sac reabsorbed and uterine cannibalism in the form of oophagy present, litter size possibly1 to 4.Adults 230 cm long or longer, size at birth at least 60 cm and possibly 78 cm. Background colour tan, reddish or yellowish todark grey-brown above and light whitish brown below, no spots or other markings as far as is known; lower eyelid strikinglylight in colour in young.Remarks: Scope of this genus follows Applegate (1974), Bass, D’Aubrey and Kistnasamy (1975c), Compagno (1984), andDingerkus (1986), in recognizing a single living species. There are also several fossil species of Nebrius (Cappetta, 1987).Compagno (1984) included Nebrius concolor and its synonyms in the synonymy of N. ferrugineus. This was accepted bysome subsequent researchers including Last and Stevens (1994) though Dingerkus (1986) recognized N. concolor as theonly valid species in this genus. Translation of Lesson’s (1830) short and unillustrated description of his Scylliumferrugineum suggests that this synonymy is correct, and that Lesson was the first scientific discoverer of the Indo-Pacifictawny nurse shark. There are only two other orectoloboids known from Waigiu, one of which Lesson described as Scylliummalaianum (= Hemiscyllium freycineti), and the other is Eucrossorhinus dasypogon. Lesson described S. ferrugineumfrom a single specimen of “roussette” (French term for scyliorhinid catsharks and catshark-like orectoloboids) that wasrusty-fawn without markings above, rosy white below, and had a white eye (the conjunctiva and upper and lower eyelids ofthe tawny nurse shark are conspicuously white, though the iris is black), had a conical short head, anterior mouth andnostrils, long barbels, large rounded labial folds at mouth, triangular teeth (exact shape not stated), body tapering from thehead, a large, triangular first dorsal fin slightly behind midlength of body, smaller triangular second dorsal, large elevatedbroad caudal fin, lobate pectoral fin, quadrilateral pelvic fins (with claspers), a lobate sharp-apiced anal fin, and wasrelatively large (137 cm [54 in] if tail length is exclusive of body length). These characters taken in combination fit the tawnynurse shark and eliminate other Indo-Australian orectoloboids including Hemiscylliidae, Parascylliidae, Stegostomatidae,Orectolobidae and Rhincodontidae. Although Lesson’s description is not as specific and detailed as Rüppell’s (1837)account and illustration of Nebrius concolor from the Red Sea, it is adequate to validate the earliest name for the tawnynurse shark as Scyllium ferrugineum.Nebrius ferrugineus and N. concolor were often retained as separate species and sometimes placed in different genera asGinglymostoma ferrugineum and Nebrius concolor. Compagno (1984) synonymized the two species and suggested thatthe dentitional differences used to separate them (Garman, 1913; Fowler, 1941) may be due to growth changes in a singlespecies (ontogenic heterodonty). The supposed differences were more compressed, more low-cusped teeth in N. concolorand less compressed, more high-cusped teeth in N. ferrugineus. At least in the material of Nebrius examined by the writerfrom the Gulf of Thailand, Philippines, and elsewhere, larger specimens over 2 m long have teeth of the N. ferrugineus type,while specimens about a metre long or less have teeth of the N. concolor type. Teeth of a specimen 1.8 m long pictured byBass, D’Aubrey and Kistnasamy (1975c) are roughly intermediate. Growth changes apparently include increase in size ofcusps relative to cusplets, shortening and broadening of the labial flange (basal ledge), and thickening and broadening ofthe teeth relative to their height.Nebrius ferrugineus (Lesson, 1830) Fig. 161Scyllium ferrugineum Lesson, 1830, Voy. aut. monde corv. La Coquille, 2(1): 95. Holotype: Specimen with body 101.5 cm(3 ft 4 in) and tail 35.5 cm (14 in) mentioned, possibly not saved. Type locality, Port Praslin, New Ireland, and Baie d’Offack,Waigiou (Waigeu), New Guinea.Synonyms: Nebrius concolor Rüppell, 1837: 62, pl. 17, fig. 2. Lectotype: Naturmuseums Senckenberg, Frankfurt,SMF-3583, 85 cm TL stuffed specimen, Massaua, Red Sea. Ginglymostoma rueppelli Bleeker, 1852: 91. Apparently anunnecessary replacement name for Nebrius concolor Rüppell, 1837, as latter is listed as a synonym. Holotype: Specimenmentioned by Bleeker is a male 1.5 m (730 lines) TL, probably from Singapore. Holotype listed as Rikjsmuseum vanNatuurlijke Histoire, Leiden, RMNH 7400, by Eschmeyer (1998: CD-ROM). Ginglymostoma muelleri Günther, 1870: 408.Apparently based on Ginglymostoma concolor Müller and Henle, 1838d: 22, pl. 6, thought by Günther to be distinct fromNebrius concolor Rüppell, 1837. Syntypes: Two stuffed specimens mentioned by Müller and Henle, in the Institut fürSpezielle Zoologie und Zoologisches Museum, der Humboldt Universität, Berlin, but possibly not extant, India. No typesmentioned by Eschmeyer (1998: CD-ROM). Scymnus porosus Hemprich and Ehrenberg, in Klunzinger, 1871: 670 (nameonly, in synonymy of Ginglymostoma muelleri). Also Hemprich and Ehrenberg, 1899: 8, pl. 6, fig. 3, Red Sea; possibly notype material, see Paepke and Schmidt (1988: 162). Nebrodes macrurus Garman, 1913: 58, pl. 8, figs 7-10. Holotype:Museum of Comparative Zoology, Harvard, MCZ-820-S, 80 cm (31.5 in) TL immature male, Port Louis Harbor, Mauritius,

Sharks of the World, Vol. 2 197according to Hartel and Dingerkus (1997: xxxvii-xxxviii). Nebrodes concolor ogilbyi Whitley, 1934: 183, fig. 1. Holotype:Queensland Museum I.1216, 955 mm female, Darnley Island, Queensland, Australia. Type status confirmed by Eschmeyer(1998: CD-ROM). Nebrius doldi Smith, 1953a: 512, fig. 30A; Smith, 1953b: 8, pls. Types: Drawing by M.M. Smith based onphotographs of a 2.9 m (9.5 ft) female? specimen from south of Beira, Mozambique, apparently no type material.Other Combinations: Ginglymostoma ferrugineum (Lesson, 1830), Ginglymostoma concolor (Rüppell, 1837).FAO Names: En - Tawny nurse shark; Fr - Requin nourrice fauve; Sp - Gata nodriza atezada.TOOTHFig. 161 Nebrius ferrugineusUNDERSIDE OF HEADField Marks: Moderately long barbels, nasoral grooves present but no circumnarial grooves, mouth well in front of eyes,eyes and gill openings lateral, spiracles minute, precaudal tail shorter than head and body, two spineless, angular dorsal finsand an anal fin, first dorsal fin much larger than second dorsal and anal fins, first dorsal-fin base over pelvic-fin bases,pectoral fins falcate, caudal fin moderately long, over one-fourth of total length. Colour: colour brown, from tan to rufous oryellowish to dark grey-brown according to habitat, and slowly changeable by the individual.Diagnostic Features: See genus Nebrius above.Distribution: Wide-ranging in the Indo-West and Central Pacific: South Africa (KwaZulu-Natal), Mozambique, Mauritius,Seychelles, Chagos Archipelago and Madagascar to Red Sea, Maldives, Persian Gulf, India, Malaysia, Indonesia,Singapore, Thailand, Viet Nam, China, Taiwan (Province of China), Japan (Ryukyu Islands, southern Honshu), Philippines(Luzon), Papua New Guinea, Australia (Western Australia, Northern Territory, and Queensland), New Caledonia, NewIreland, Samoa, Palau, Marshall Islands, and Tahiti.Habitat: A large, tropical inshore shark of the continental and insular shelves of the Indo-Pacific, often in the intertidal inwater scarcely able to cover it and from the surf line down to a few metres depth, commonly at 5 to 30 m and ranging down toat least 70 m on coral reefs. It occurs on or near the bottom in lagoons, in channels, or along the outer edges of coral and

198 FAO Species Catalogue for Fishery Purposes No. 1rocky reefs, in areas with seagrass and sand on reefs, sandy areas near reefs and off sandy beaches. It prefers shelteredareas in crevices and caves on reefs but often occurs in more exposed areas in depressions or crevices. Young prefercrevices in shallow lagoons but adults are more wide-ranging.Biology: The tawny nurse shark is primarily nocturnal, resting in the daytime in shelters but prowling slowly about aroundreefs at night, although some individuals may be active in the day. In Madagascar it is described as day and night-active, andin captivity they get quite active and vigorous when food is presented to them during the day. They are social, gregarioussharks when at rest and form resting aggregations of two to a half-dozen or more in shelters, and are often seen piled inertlyacross or on top of one another. When resting, they are extremely sluggish. This shark has a limited home range, andindividuals often return to the same area every day after foraging.Reproduction ovoviviparous (aplacental viviparous) with cannibal vivipary or uterine cannibalism in the form of oophagy.This shark has been described as an oviparous or post-oviparous shark that retains the egg-cases until they hatch and theyoung are born, but recent evidence indicates that this is incorrect. Pregnant females collected from Okinawa have had oneor two foetuses per uterus, 297 to 595 mm (the latter near term), with the yolk sac reabsorbed and a greatly expandedstomach filled with yolky material in foetuses 338 and 595 mm long, and also had cased eggs in the uterus. Apparently thisspecies practices oophagy on relatively large, cased nutritive eggs (unlike many lamnoids which have very small nutritiveeggs), and is the first orectoloboid known to have uterine cannibalism. It is not known whether foetuses of this species eatother foetuses (adelphophagy) as with the sand tiger (Carcharias taurus). The presence of two foetuses 338 and 297 mm inthe same uterus, with the smaller one slender and the larger bloated with yolk, suggests that competition between siblingsfor the relatively big eggs is likely and could even eliminate the less successful sibling. Adelphophagy is less likely althoughearly stages need to be examined to eliminate it. Number of young per litter uncertain; at least four young per uterus hasbeen suggested from cased eggs, but the size and enormous girth of the near-term Okinawan foetus and the two smallerfoetuses in a litter suggests that litters are smaller, possibly one or two per uterus or even one per female, and that numbersof cased eggs in the uteri cannot be used to extrapolate litter sizes in this species. In captivity adult females lay cased eggson the bottom, but these do not develop, and could be nutritive and unfertilized. Such free eggs may have been the basis ofthe suggestion that this species is oviparous. The tawny nurse shark breeds in July and August off Madagascar.Food of this shark includes corals, crabs, lobsters and other crustaceans, octopi, squid and probably other cephalopods, seaurchins, and reef fish including surgeonfish (Acanthuridae), queenfish (Carangidae) and rabbitfish (Siganidae), andoccasionally sea snakes. While foraging the tawny shark moves along the bottom and explores depressions, holes andcrevices in reefs. When it detects prey it places its small mouth very close to the victim, and uses its large pharynx as apowerful suction pump to rapidly suck in reef organisms that may be out of reach of its teeth. A few large individualsdissected by the writer had quantities of small, active reef fishes in their stomachs, presumably sucked in by the sharks asthe prey fishes lay inert in shelters or on the bottom at night. Individuals caught by fishermen may reverse this sucking action,and blast streams of water out of their mouths and into the faces of their captors; they are said to make a grunting soundbetween blasts. It is not known if spitting water is deliberate and defensive or if the sharks are actually aiming the water at theanglers. They also tend to spin when hooked on a line, making them difficult to handle and subdue.The body form of the tawny nurse shark (littoral morphotype) is more fusiform and streamlined than other nurse sharks, withnarrow-based, falcate, plesodic pectorals, pointed dorsal and anal fins with the anal-fin apex raked posterior to the free reartip, a short ventral caudal-fin lobe, lateral eyes and gills, a narrow head, flat wedge-shaped snout, and compressed,semi-blade-like, imbricate teeth in discrete series. The tawny nurse shark superficially resembles certain other large, partlyor mostly sympatric, active reef sharks including the sand tiger shark (Carcharias taurus), sicklefin lemon shark(Negaprion acutidens), and reef whitetip shark (Triaenodon obesus). Whether it is behaviourally divergent from other nursesharks awaits a detailed comparative study of nurse shark behaviour. Its status as a game fish in Australia, unlike the nurseshark in the western Atlantic, suggests that it may be a more active swimmer when not resting on the bottom.Size: Maximum about 314 to 320 cm, though most individuals are smaller; size at birth has been reported as about 40 cm(Fourmanoir and Laboute, 1976) but subsequent data from a pregnant female captured off Okinawa suggest that it mayreach 60 cm or more at birth (Teshima et al. 1995), while a 79 cm female from Navotas market in Manila, Philippines,examined by the writer, has a somewhat bloated stomach full of yolk, suggesting that it was newborn or a term foetus; malesare mature at about 250 cm and reach at least 301 cm; adult females are 230 to at least 290 cm.Interest to Fisheries and Human Impact: Common or formerly common in areas where it occurs, and caught inshore byfishermen in Pakistan, India, Thailand, and Philippines, and probably widely captured elsewhere. It is utilized fresh anddried-salted for human food, its liver is rendered for oil and vitamins, its fins are used in the oriental sharkfin trade, and offal isprocessed into fishmeal. Its thick, armour-like hide is potentially valuable for leather. Off Queensland, Australia, it has beenfished as a big-game shark, and large individuals are prized as powerful fighters by sports anglers. Apart from anglers whotarget this fish, it is apparently primarily caught only as an untargeted bycatch of fisheries in inshore waters in nets, on linegear, and in fish traps.This has been described as a much more docile species than its close relative, Ginglymostoma cirratum, and apparentlytolerates close proximity of divers and usually allows humans to touch and play with it without biting. However, there are afew records of these sharks biting their tormentors, and clamping tightly onto them. Because of its size, strength, powerfuljaws and small but sharp cutting teeth, the tawny shark should be treated with the respect due it. It is a favourite species forobservation by ecotouristic divers within its enormous range, and dive sites where it is viewed are known from Thailand(Andaman Sea) and the Solomon Islands.

Sharks of the World, Vol. 2 199This is a tough, hardy shark that readily survives in captivity and makes an excellent and handsome display animal. It is keptin aquaria in Europe, Japan (Okinawa), Singapore, and the United States. In Singapore in 1996 the writer saw several largeNebrius about 2 m long being hand-fed by a SCUBA diver in a big oceanarium tank. The sharks piled on top of theirbenefactor and hid him from sight while eagerly scrambling to obtain food.The conservation status of this shark is uncertain and urgently needs investigation despite its wide range. In some areas,including the Gulf of Thailand where it was commonly caught in the 1960s, it may have been depleted due to increasingfisheries activity and habitat degradation. Also, reef habitats have been extensively damaged or destroyed by dynamitingand poisoning in parts of its range, including Indonesia and Philippines, which probably have had an adverse effect on thisspecies both directly and through decimation or elimination of its prey. Its docility and inshore habitat makes it particularlysusceptible to a wide variety of fishing gear, to harassment and injuries by divers, and to reef destruction and pollution.Local Names: Tawny shark, Spitting shark, Rusty shark, Rusty catshark, Sleepy shark, Madame X (Australia); Giant sleepyshark (South Africa); Roussette ferrugineuse (French); Yahiya (Madagascar); Endormi (Seychelles); Kalouyon (Guébéens);Koumoune (Waigiu); Be kal mora (Sinhalese); Ô-tenjikuzame (Japan).Remarks: Several individuals from Okinawa and Honshu, Japan and at least one from Taiwan (Province of China) havebeen found with their second dorsals missing or rudimentary. It is not known why this occurs in these localities, but there aresome interesting possibilities that might be investigated including teratogenic effects of chemical pollutants.Literature: Lesson (1830); Rüppell (1837); Müller and Henle (1839); Gray (1851); Dumeril (1865); Günther (1870); Jordanand Seale (1906); Garman (1913); Whitley (1934, 1940); Fowler (1941); Misra (1947); Bigelow and Schroeder (1948);Fourmanoir (1961); Gohar and Mazhar (1964); Marshall (1965); Applegate (1974); Bass, D’Aubrey and Kistnasamy(1975c); Fourmanoir and Laboute (1976); Johnson (1978); Randall (1980); Yoshino et al. (1981); Uchida (1982); Compagno(1984, 1988); Nakaya and Shirai (1984); Bass (1986); Dingerkus (1986); Taniuchi and Yanagisawa (1987); Compagno,Ebert and Smale (1989); Anderson and Ahmed (1993); Michael (1993); Seret (1994); Last and Stevens (1994); Teshima etal. (1995, 1999).Pseudoginglymostoma Dingerkus, 1986Genus: Pseudoginglymostoma Dingerkus, 1986, Proc. 2nd. Int. Conf. Indo-Pacific Fishes, Tokyo: 240.Type Species: Ginglymostoma brevicaudatum Günther, in Playfair and Günther, 1866, by original designation.Number of Recognized Species: 1.Synonyms: None.Diagnostic Features: Head in dorsal or ventral view very broad and parabolic in adults. Snout bluntly rounded in lateralview, short and with preoral length about 34.5 to 41.6% of mouth width. Eyes very small and 0.8 to 1.0% of total length,situated dorsolateral on head and with strong subocular ridges below them; ventral edges of eyes just above level of dorsalends of gill slits. Gill openings dorsolaterally situated on head and not visible from below, just reaching horizontal head rimfrom above. Nostrils nearly terminal on snout. Nasal barbels very short, stubby, not very tapered, less than 1% of totallength, and not reaching mouth. Lower lip not trilobate and without orolabial grooves. Distance between lower labial furrowsabout 1.2 to 1.4 times their length. Tooth rows 24 to 27/22 to 27 (adults), functional series at least 3 to 4/4 to 5. Teeth notcompressed and not imbricated, functional series not separated from replacement series by toothless space. Tooth crownfeet very narrow, cusps large, cusplets very small and one or two on each side; teeth with moderately broad basal ledges.Teeth orthodont and with a central pulp cavity. Body cylindrical. Lateral trunk denticles teardrop-shaped and elongated inadults, with a low medial ridge, lateral ridges weak or absent, and with a fairly narrow pointed cusp. Pectoral fins broad andnot falcate in adults, apically rounded. Pectoral-fin origins slightly in front of third gill slits. Pectoral fins semiplesodic and withradials reaching only 55% into fin web, radial segments four. Pelvic fins rounded in adults. Dorsal fins apically rounded. Firstdorsal-fin origin over or slightly behind pelvic-fin midbases. Second dorsal fin about as large as first dorsal fin. Anal fin aboutas large as first dorsal fin, with rounded apex. Anal-fin apex about under anal free rear tip, origin about opposite seconddorsal-fin origin, posterior margin ends in front of lower caudal-fin origin. Caudal fin short with dorsal caudal-fin margin lessthan 25% of total length (adults); caudal fin broad and deep with depth 40 to 45% of dorsal caudal-fin margin; no ventralcaudal-fin lobe in adults; preventral caudal-fin margin about as long as postventral margin and 80 to 100% of it; terminalcaudal-fin lobe moderate and 22.2 to 26.3% of dorsal caudal-fin margin. Total vertebral count 135 to 143, monospondylousprecaudal count 35 to 37, diplospondylous caudal count 49 to 54 and 36 to 39% of total count. Jaws broadly arcuate.Intestinal valve count 15. Mode of reproduction and litter size unknown. Adults 59 to 75 cm long, size at birth or hatchingunknown but probably less than 30 cm. Colour: background colour dark brown above and slightly lighter below butnoticeably dark on both surfaces; no spots or other markings in adults but colour of young unknown; lower and upper eyelidsdark in adults.Remarks: Scope of this genus follows Dingerkus (1986), with a single living species, Pseudoginglymostomabrevicaudatum.

200 FAO Species Catalogue for Fishery Purposes No. 1Pseudoginglymostoma brevicaudatum (Günther, 1866) Fig. 162Ginglymostoma brevicaudatum Günther, in Playfair and Günther, 1866, Fish. Zanzibar, London: 141, pl. 21. Holotype:British Museum (Natural History), BMNH-1867.3.9.423, stuffed dried adolescent male about 590 mm TL, length in originalaccount 640 mm, Zanzibar.Synonyms: None.Other Combinations: None.FAO Names: En - Short-tail nurse shark; Fr - Requin-nourrice à queue courte; Sp - Gata nodriza rabicorta.TOOTHFig. 162 Pseudoginglymostoma brevicaudatumUNDERSIDE OF HEADField Marks: Very short barbels, nasoral grooves present but no circumnarial grooves; eyes and gill openings dorsolateral,mouth well in front of eyes, spiracles minute, precaudal tail shorter than head and body, two spineless, broadly rounded,equal-sized dorsal fins and an equally large anal fin, caudal fin short, less than one-fourth of total length, colour dark brown,without spots or other markings.Diagnostic Features: See genus Pseudoginglymostoma above.Distribution: Confined to the western Indian Ocean offTanzania, Kenya, Madagascar and possibly Mauritiusand the Seychelles.Habitat: This little-known inshore bottom shark occurson the continental and insular shelves of East Africa andMadagascar, but details are lacking on its habitat exceptthat it occurs on coral reefs. Depth data are not available.Biology: The biology of the short-tail nurse shark,including breeding and feeding habits, is poorly known.An adult or near-adult female had large nidamentalglands, suggesting that the species may be egg-layingbut this needs confirmation (see below). It also is notknown if this species is social and congregates in groupsas with the larger nurse sharks. It is reported to survivefor several hours out of water. Abundant off East Africa afew decades ago (Bass, D’Aubrey and Kistnasamy,1975c), but its current status is uncertain.A female short-tailed nurse shark has lived over 33 yearsand a male three years in captivity at the Artis Zoo, Amsterdam. These are under study by A. Dral, E. Bruins and P. Bor (pers.comm.), who are preparing a detailed account for publication. Both individuals are alive at the time of writing and the femalehas laid infertile eggs in strong egg cases two at a time for the past ten years. This suggests that the short-tailed nurse sharkis oviparous but needs confirmation. These sharks mate in captivity but so far no eggs have been hatched as the pair eattheir eggs. These sharks are slow-moving, nocturnally active, sit on the open bottom or hide in holes or crevices during theday, and will only move during the day when fed. They eat annelid worms, raw and cooked mussels, cut raw fish andshrimps; presumably the species eats small fishes, molluscs and crustaceans in the wild. The two sharks arenon-aggressive in community tanks and the female has been kept with much larger nurse sharks (Ginglymostomacirratum) without incident.

Sharks of the World, Vol. 2 201Size: Maximum at least 75 cm; two adult males examined were 59 to 75 cm while a live captive adult male (Artis Zoo,Amsterdam) is 65 cm long; a female examined was immature at 52 cm while another was adolescent at 56 cm and a thirdwas adult or nearly so (with large oviducts and nidamental glands but without oviducal eggs or foetuses) at 53 cm. A captiveadult female at the Artis Zoo is 70 cm long.Interest to Fisheries and Human Impact: Interest to fisheries probably limited, apparently fished locally in artisanalfisheries and landed as a bycatch of other fisheries. The skin of this shark is exceptionally tough, as in other nurse sharks,and is possibly of use for leather. It is readily amenable to captivity and grows to a more reasonable maximum size forsmaller aquaria than Nebrius ferrugineus or Ginglymostoma cirratum, but it’s status in the aquarium trade is uncertain. Itapparently is seldom kept in public aquaria. The conservation status of this interesting and distinctive little shark is unknownbut is of concern as it has a limited distribution in inshore tropical waters of East Africa and occurs in some areas thatcurrently support heavy inshore fisheries. It could be adversely affected by overfishing and destruction of coral reefs.Local Names: Nurse shark, Shorttail nurse shark, Papa isengezi or Sleepy shark (Zanzibar), Endormi (Seychelles).Remarks: The writer examined the holotype in the British Museum (Natural History) and five other specimens in thecollections of the J.L.B. Smith Institute of Ichthyology, Grahamstown, South Africa.Literature: Günther in Playfair and Günther (1866); Günther (1870); Garman (1913); Fowler (1941, 1967a); Smith andSmith (1963); Bass, D’Aubrey and Kistnasamy (1975c); Compagno (1984, 1988); Dingerkus (1986); Bass (1986); AntonDral, Eugene Bruins and Peter Bor (pers. comm.).2.3.7 Family RHINCODONTIDAEFamily: Rhinodontes Müller and Henle, 1839, Syst. Beschr. Plagiost., pt. 2: 77. Placed on the Official List of Family-GroupNames in Zoology (Name no. 559) by the International Commission on Zoological Nomenclature, 1984, Opinion 1278, Bull.Zool. Nomen., 41(4): 215 as the emended Family Rhincodontidae.Type Genus: Rhincodon Smith, 1829.Number of Recognized Genera: 1.Synonyms: Subfamily Rhinodontini Bonaparte, 1846, v. 3 (no pagination); Tribe Rhineodontiana (Family Squalidae) Gray,1851: 66. Family Rhineodontis “Müll. and Henle” Gray (1851: 66). Family Rhinodontoidei Bleeker, 1859: XII. FamilyRhinodontoidae Owen, in Gill, 1862b: 397. Family Rhinodontidae Günther, 1870: 396. Family Rhineodontidae Jordan andFowler, 1903: 626. Subfamily Rhinodontinae (Family Lamnidae) Goodrich in Lankester, 1909: 150. Family RhincodontidaeGarman, 1913: 12, 41. Type genus: Rhincodon Smith, 1829. Subfamily Rhineodontini (Family Orectolobidae) Berg, 1940:380. Subfamily Rhincodontinae (Family Rhincodontidae) Fowler, 1947: 10. Family Rhiniodontidae Compagno, 1973: 28.Type genus: Rhiniodon Smith, 1828, emendation of Family Rhinodontes Müller and Henle, 1839, used by Compagno(1984: 209).FAO Names: En - Whale sharks; Fr - Requins baleine; Sp - Tiburones ballena.Diagnostic Features: Head very broad and greatly flattened, without lateral flaps of skin. Snout truncated. Eyes laterallysituated on head and without subocular ridges below them. Eyes without movable upper eyelids or subocular pockets andridges. Spiracles moderate-sized but much smaller than eyes, without raised external rims; spiracles behind but not beloweyes. Gill slits very large, fifth gill slit well-separated from fourth and not overlapping it; internal gill slits with unique filterscreens, consisting of transverse lamellae that cross each gill slit, with ramose processes on their inner surfaces thatinterconnect to form the filters. Nostrils with rudimentary barbels and no circumnarial folds and grooves. Nasoral groovesvery short and shallow. Mouth very large, transverse and terminal on head. Lower lip not trilobate and without lateralorolabial grooves connecting edge of lip with medial ends of lower labial furrows, without a longitudinal symphysial grooveon chin. Lower labial furrows ending far lateral to symphysis, not connected medially by a mental groove or groove and flap.Teeth not strongly differentiated in upper and lower jaws, with symphysial teeth not enlarged and fang-like. Tooth row countextremely high, in over 300 rows in either jaw of adults and subadults. Teeth with a strong medial cusp, no cusplets and nolabial root lobes. Teeth osteodont with crown partially filled with a plug of osteodentine. Body cylindrical or moderatelydepressed, with prominent ridges on sides. Precaudal tail shorter than body. Caudal peduncle with strong lateral keels andan upper precaudal pit. Pectoral fins very large, relatively narrow and falcate. Pectoral fins plesodic and with fin radialsstrongly expanded into fin web. Pectoral propterygium small and fused to mesopterygium; pectoral-fin radial segments 3 to10, and with longest distal segments about 0.8 times the length of longest proximal segments. Pelvic fins much smaller thanfirst dorsal fin but subequal to second dorsal and anal fins, much smaller than pectoral fins and with anterior margins about0.3 times the pectoral-fin anterior margins. Claspers without mesospurs, claws or dactyls. First dorsal fin much larger thansecond. First dorsal-fin origin well anterior to pelvic-fin origins and over abdomen behind pectoral-fin free rear tips, firstdorsal-fin insertion over the pelvic-fin bases. Anal fin about as large as second dorsal, with broad base, angular apex, origin

202 FAO Species Catalogue for Fishery Purposes No. 1about opposite first third of second dorsal fin base, and insertion separated by a space somewhat greater than base lengthfrom lower caudal-fin origin. Caudal fin elongated and semicrescentic, strongly heterocercal with its upper lobe at a highangle above the body axis (lower in young than adults and subadults); dorsal caudal-fin margin less than a third as long asthe entire shark. Caudal fin with a vestigial terminal lobe and subterminal notch but with a strong ventral lobe (longer in adultsthan young), preventral and postventral margins strongly differentiated and deeply notched. Vertebral centra withwell-developed radii and prominent annuli connecting them. Total vertebral count 174, monospondylous precaudal count42, diplospondylous precaudal count 40, diplospondylous caudal count 92, and precaudal count 82. Cranium very broadand greatly expanded laterally. Medial rostral cartilage rudimentary, reduced to a low nubbin. Nasal capsules greatlydepressed, slightly fenestrated anteriorly, internarial septum low, broad and depressed. Orbits with enlarged fenestrae forpreorbital canals, medial walls not fenestrated around the optic nerve foramina although foramen itself is very large.Supraorbital crests present on cranium and laterally expanded and pedicellate. Suborbital shelves very broad and notgreatly reduced. Cranial roof with a continuous fenestra from the anterior fontanelle to the parietal fossa. Basal plate ofcranium with pairs of separate carotid and stapedial foramina. Adductor mandibulae muscles of jaws with three divisions.Preorbitalis muscles extending onto posterodorsal surface of cranium. No anterodorsal palpebral depressor,rostromandibular, rostronuchal or ethmonuchal muscles. Valvular intestine of ring type with 69 to 74 turns. Developmentovoviviparous. Size gigantic with adults 700 cm or longer, while young are born at about 55 to 64 cm. Colour pattern unique,consisting of small to large white or yellowish spots and vertical and horizontal stripes in the form of a checkerboard on adark background.Local Names: Whale sharks, Ching sha k’o, Jimbeizame-ka (Japan); Kitovye akuly (Russia).Remarks: As recognized here the family Rhincodontidae includes only a single genus and species, the whale shark,Rhincodon typus. The genus and species were first described as Rhiniodon typus by Smith (1828) in a Cape Townnewspaper from a specimen stranded in Table Bay, South Africa. A separate family Rhinodontes was later proposed for it byMüller and Henle (1839) who used a variant spelling Rhinodon for the genus. There has been considerable variation inspelling of the generic name (Rhinodon, Rhineodon, Rhincodon, Rhiniodon, Rhinecodon, Rhinocodon, Rineodon andRhinchodon) and by extension the family name of the whale shark (Rhinodontidae, Rhincodontidae, Rhineodontidae andRhiniodontidae), and much usage of certain of the variants. Andrew Smith, the describer of the whale shark, and JohannesMüller and Friedrich Henle, the describers of a family for it, used several variants on the genus, which caused confusion forsubsequent authors. Although the original Rhiniodon has priority, the variants Rhincodon and particularly Rhineodon andRhinodon have had far more usage over the last two centuries.Following Bigelow and Schroeder (1948) the variant Rhincodon has received considerable use, and proposals to stabilize it(Robins and Lea, 1975; Swift, 1977; Wheeler, 1982) were presented to the International Commission on ZoologicalNomenclature. Brooke and Bass (1976) supported the earliest published spelling, Rhiniodon, although noting thatRhiniodon Smith, 1828 was a possible misspelling that Andrew Smith had hand-corrected to Rineodon on a bound copy ofhis paper (Smith, 1828). Rineodon was also used by Müller and Henle (1838a), possibly on the advise of Andrew Smith, butlater changed to Rhinodon (Müller and Henle, 1839). Hubbs, Compagno and Follett (1976) proposed that the earliestspelling, Rhiniodon should be preserved for the whale shark because of priority, more correct orthography than Rhincodon,and because the use of Rhincodon has not been universal since Bigelow and Schroeder’s work (and little-used prior to itcompared to Rhinodon and particularly Rhineodon). Compagno (1984) preferred to use the earliest spelling (Rhiniodonand emended family name Rhiniodontidae) but while that work was in press the International Commission on ZoologicalNomenclature (1984, Opinion 1278) stabilized Rhincodon and Rhincodontidae.Dingerkus (1986) included the genera Pseudoginglymostoma, Ginglymostoma, Nebrius and Stegostoma in this family, butthe original arrangement of Rhincodon alone in Rhincodontidae is followed here on the basis of the extremely derivedmorphology of the whale shark, uncertain relationships with other ‘higher’ orectoloboids, and pending further studies onorectoloboid phyletics.Rhincodon Smith, 1829Genus: Rhincodon Smith, 1829, Zool. J., 4: 443. Placed on the Official List of Generic Names in Zoology (Name no. 2219,International Commission on Zoological Nomenclature, 1984, Bull. Zool. Nomen., 41(4): 215). Rhincodon was used twice inthe 1829 publication.Type Species: Rhiniodon typus Smith, 1828, as interpreted by the International Commission on Zoological Nomenclature,1984, Opinion 1278, Bull. Zool. Nomen., 41(4): 215.Number of Recognized Species: 1.Synonyms: Genus Rhiniodon Smith, 1828: 2. Type species: Rhiniodon typus Smith, 1828, by monotypy and by use of thespecies name typus. Placed on the Official Index of Rejected and Invalid Generic Names in Zoology (Name no. 2141) by theInternational Commission on Zoological Nomenclature (1984, Opinion 1278: 215). Rhiniodon was twice cited in the originaldescription, but possibly was a typographical error or correction by the printer (Brooke and Bass, 1976: 5). GenusRhinchodon Smith, 1829: 534 (index). Variant spelling. Genus Rineodon Müller and Henle, 1838a: 37. Also Swainson,1838: 142. Variant spelling. Genus Rhineodon Müller and Henle, 1838c: 84. Variant spelling. Genus Rhinodon Müller and

Sharks of the World, Vol. 2 203Henle, 1839: 77. Variant spelling. Genus Rhiniodon Swainson, 1839: 317. Variant spelling. Genus Rhinecodon Agassiz,1845: 55. Variant spelling. Genus Rhinodon Smith, 1849: pl. 26 and accompanying text, pages not numbered; Smith usedRhinodon in this extended account and illustration of the whale shark rather than his earlier Rhiniodon or Rhincodon.Variant or emended spelling. Genus Micristodus Gill, 1865: 177. Type species: Micristodus punctatus Gill, 1865, bymonotypy. Genus Rhinocodon International Commission on Zoological Nomenclature, 1984: 216). Attributed to Herald,1961, but examination of the writer’s copy indicated uniform usage of Rhincodon in the text (p. 22) and index (p. 302),suggesting that the variant spelling if correctly cited was a possible typographical error in a different edition of Herald’s book.Diagnostic Features: See family Rhincodontidae above.Rhincodon typus (Smith, 1828) Fig. 163Rhiniodon typus Smith, 1828, S. African Comm. Adv., 3(145): 2. Holotype: Museum National d’Histoire Naturelle, Paris,MNHN 9855, 4 600 mm stuffed, mounted male specimen, Table Bay, South Africa. The specific name typus Smith, 1828 wasplaced on the Official List of Species Names in Zoology (Name no. 2901) by the International Commission on ZoologicalNomenclature, 1984, Opinion 1278, Bull. Zool. Nomen., 41(4): 215.Synonyms: Rhincodon typus Smith, 1829: 443. Rhinodon typicus Müller and Henle, 1839: 77, pl. 35. Variant or emendedspelling of Rhincodon typus. This spelling was also used by Smith, 1849: pl. 26 and accompanying text, pages notnumbered, in his extended description of the whale shark. Micristodus punctatus Gill, 1865: 177. Holotype: US NationalMuseum of Natural History, USNM-27234, renumbered as USNM-232756, a pair of dried dental band halves, Gulf ofCalifornia. Status of holotype from Howe and Springer (1993: 12) and by the writer’s examination of the specimen.Rhinodon pentalineatus Kishinouye, 1901: 694, figs 1-2. Holotype: Specimen originally in private collection of T. Oseko,from Cape Inubo, Japan, whereabouts unknown according to Eschmeyer (1998: CD-ROM).Other Combinations: Rhineodon typus (Smith, 1828), Rhineodon typicus (Müller and Henle, 1839).FAO Names: En - Whale shark; Fr - Requin baleine; Sp - Tiburón ballena.Fig. 163 Rhincodon typusField Marks: An unmistakable huge shark, one of three large filter-feeding species (apart fromdevel rays), with a broad, flat head and truncated snout, immense transverse and virtuallyterminal mouth in front of eyes, minute, extremely numerous teeth, and unique filter screens onits internal gill slits; prominent ridges on sides of body with the lowermost one expanding into aprominent keel on each side of the caudal peduncle, a large first dorsal and small second dorsaland anal fin, lunate or semilunate caudal fin without a prominent subterminal notch. Colour: aunique checkerboard pattern of white or yellow spots, horizontal and vertical stripes on a grey,bluish, reddish or greenish brown dorsal surface, abruptly white or yellowish on the underside ofthe body.LATERAL VIEW OF HEADDiagnostic Features: See family Rhincodontidae above.Distribution: Circumglobal in all tropical and warm temperate seas, oceanic and coastal. Western Atlantic: Canada (southof Nova Scotia), United States (Gulf of Maine and New York south to North Carolina and Florida, and Gulf of Mexico coastfrom Florida to Texas), Mexico (Gulf of Mexico coast to Quintana Roo), Belize, Honduras, Panama, Colombia, Venezuela,and central Brazil, also Bermuda, Bahamas, Caribbean including Cuba and Haiti. Eastern Atlantic: Azores, Senegal,Mauritania, Cape Verde Islands, Côte d’Ivoire, Gulf of Guinea, Gabon, Angola, South Africa (Western Cape Province).Indo-West and Central Pacific: East coast of South Africa (Western Cape, Eastern Cape, KwaZulu-Natal), Mozambique,Madagascar, Seychelles, Kenya, Tanzania, Somalia, the Red Sea and Straits of Bab-al-Mandab (Eritrea, Sudan, SaudiArabia), Yemen, Oman, Persian Gulf, the Maldives, Pakistan, India (both coasts), Sri Lanka, possibly Bangladesh, Malaysia(including Sabah, Borneo), Singapore, Thailand (including Gulf of Thailand), Viet Nam, China, Taiwan (Province of China),

204 FAO Species Catalogue for Fishery Purposes No. 1Japan (Southeastern Honshu, Okinawa, and Bonin Islands), Philippines, Indonesia (Kalimantan, Borneo; Java), NewGuinea (Papua-New Guinea and Irian Jaya, Indonesia), Australia (Western Australia, Queensland, Northern Territory, withisolated records from New South Wales and Victoria), New Caledonia, Hawaiian Islands, Tuomotu Archipelago, PhoenixIslands. Eastern Pacific: United States (southern California), Mexico (Baja California and Gulf of California south toAcapulco, Oaxaca, and Chiapas), El Salvador, Nicaragua, Costa Rica, Panama, Ecuador, Peru, and northern Chile(including open ocean from west of Cabo San Lucas to the Galapagos Islands, and between Panama and the HawaiianIslands).Habitat: An epipelagic and neritic, oceanic and coastal, tropical and warm-temperate pelagic shark, often seen far offshorebut regularly coming close inshore off beaches and coral reefs and sometimes entering lagoons of coral atolls. In thewestern Pacific it apparently prefers areas where the surface temperature is 21 to 25°C with cold water of 17°C or lessupwelling into it, and salinity of 34.0 to 34.5 ppt. Recent satellite tagging (Eckert and Stewart, 1996) in the Gulf of Californiasuggested that the sharks prefer water over 26°C and up to 34°C, although they were recorded in water masses attemperatures down to 10°C; the sharks tended to move out of parts of the Gulf of California where surface water cooledbelow 26°C with upwelling. These conditions may be optimal for production of plankton and small to moderate-sizednektonic organisms, all of which are prey of the whale shark. The whale shark is generally seen or otherwise encounteredclose to or at the surface in warm waters, although recent satellite tagging in the Caribbean Sea off Belize shows that thewhale shark may dive to depths of 700 m and may transit in cold deep water down to 7.8°C (R. Graham, pers.comm.). OffNingaloo Reef in Western Australia a sonic-tagged shark frequently dived during two 18 hr tracks and ranged from thesurface to near the bottom at depths of 40 to 70 m (Stevens et al., 1997), with salinity at 34.9 to 35.2 ppt and temperatures of26.8 to 27.5°C at the surface to 26.4 to 25.4°C at the bottom.Strandings of whale sharks are common in some areas, including both coasts of South Africa, and it is suspected that off thewest coast of South Africa whale sharks ride the warm Agulhas current during the summer into areas where upwellingoccurs and plumes of cold bottom water hit the surface. It was thought that sharks may be stunned or even killed by suddenchilling and then wash up on the shore with no signs of disease or physical damage (Beckley et al., 1997). Tracking of whalesharks in water down to 10°C in the Gulf of California and their presence near patches of upwelling water down to 6°C(Eckert and Stewart, 1996) makes this simple explanation suspect however, and an explanation for the phenomenon ofstranding off the Western Cape requires further research. Rough seas and sudden storms along the narrow continentalshelves of KwaZulu-Natal, South Africa (which is subtropical and optimal for whale sharks) may contribute to strandings onthe beaches there.Biology: The whale shark is a facultatively social shark, and has been recorded as single individuals or in schools oraggregations of up to hundreds of sharks. In the Indian Ocean it is common around the Seychelles, Mauritius, Zanzibar,Kenya, Madagascar, Mozambique and northernmost KwaZulu-Natal. In the western Pacific it is common in the Kuroshiocurrent in the fishing grounds for skipjack (Scombridae). It is reportedly abundant in the Gulf of California and from Cabo SanLucas to Acapulco in the eastern Pacific, and in the Gulf of Mexico and the Caribbean in the western Atlantic. Aerial spottingof whale sharks has been successfully attempted off Kenya, South Africa, and Western Australia, as well as whale sharksurveys by boat in these same localities (Gifford, 1994; Wamukoya, Mirangi and Ottichilo, 1995; Beckley et al., 1997;Stevens et al. 1997; Colman, 1997), while whale shark observation records are collected routinely from dive operators inWestern Australia (as with the various basking shark sighting programs in the United Kingdom). The US Shark ResearchInstitute (http://www.sharks.org) runs an Internet-based program encouraging people who sight whale sharks to send inreports by eMail to their central database, which so far has received several hundred responses.

Sharks of the World, Vol. 2 205Whale sharks are highly migratory, with their movements probably timed with localized blooms of planktonic organisms andchanges in temperatures of water masses. Several whale sharks have been tagged with radio satellite tags using theARGOS system in the Gulf of California (Eckert and Stewart, 1996) over times of up to a year, with maximum speedrecorded about 2.3 km/hr. Satellite tracking has revealed extensive movements, and tagged whale sharks have moved outof the Gulf of California along rugged bottom topography of the Clipperton Fracture Zone to the Revilligigedo and ClippertonIslands, with one shark travelling 1 723 km in about three months and another 3 708 km in five months. Satellite taggingsuggested that whale sharks in the Gulf of California showed some segregation by size, with smaller sharks frequenting thenorthern Gulf and possibly absent from the southern part. Observation of whale sharks off Ningaloo Reef, Western Australiasuggests that mostly adolescent males between 6 and 7.5 m length are seen there, with one (possibly mature) malemeasured live with a rope at 9.1 m; between 80 and 91% of the sharks seen in 1996 and 1997 were male (Stevens et al.,1997; Gunn et al. 1999). Some 53 individuals were ‘body-printed’ with scarring and colour patterns recorded, and 29resighted from one to 14 times (Stevens et al. 1997) over at least two years. In South Africa SCUBA divers working under theauspices of the US Shark Research Institute tag whale sharks underwater using modified spearguns with detachablespearheads attached to giant spaghetti tags, and have resighted the tagged sharks later by approaching them closely andreading the tag numbers (Gifford, 1994); at least one of these sharks had been resighted off the Seychelles (ca 3 700 km).Satellite tagging has also been attempted on whale sharks off KwaZulu-Natal, South Africa, and off Ningaloo Marine Park,with tracks of limited range so far.Whale sharks are often associated with schools of pelagic fish, especially scombrids. They apparently show curiosity in thepresence of people and will approach divers and boats, possibly to examine them. The behaviour and sociobiology of thewhale shark is sketchily known at present, but as with the basking and white sharks their detailed behaviour should beamenable to elucidation by divers and observers in boats using cinematography as well as telemetry. The most extensiveclose-in behavioural research to date has been at the famous whale shark viewing site at Ningaloo Reef (Ningaloo MarinePark), Australia (Stevens et al., 1997; Gunn et al., 1999), where sonic tracking and archival tagging showed that whalesharks made numerous dives during a 24 hour period, ranging from the surface to near the bottom and varying in speed from0.1 to 1.8 m/sec. The sharks’ dives appeared to be independent of hydrographic features and seemed to be associated withfood search. Whale sharks showed an ability to accurately sense the bottom and swim very close to it without colliding.Some behaviour patterns observed in association with and in part in reaction to ecotouristic snorkel-divers at Ningalooinclude diving, where sharks descended out of view of the divers; porpoising, a movement away from the surface, but not outof sight; changes in speed, where sharks go slow (snorkelers easily keep up with the sharks at 0 to 1 knots), medium (wheredivers strain to keep up at a speed of 1 to 2 knots), and fast (where sharks pull away from the divers at over 2 knots); degreeof mouth distension (mouth closed to fully open on a six-point scale, possibly related to feeding); banking, in which the sharkrolls and presents its dorsal surface towards the recorder; and eye-rolling, which was observed by divers next to the whaleshark’s head.The mode of reproduction of the whale shark is apparently ovoviviparous, but it was long disputed and assumed to beoviparous by some authors. In 1953 a large eggcase, 30 cm long, 14 cm wide and 9 cm high containing a nearly full-term,36 cm embryo whale shark was collected from the Gulf of Mexico, and the assumption was made that the species isoviparous (Baughman, 1955; Reid, 1957; Garrick, 1964; Bass, D’Aubrey and Kistnasamy, 1975c). However, the rarity of‘free-living’ whale-shark eggs, the extreme thinness of its walls and lack of tendrils on the only known deposited eggcase,the considerable yolk and partially developed gill sieve in the embryo within it, and the presence of umbilical scars on largerfreeliving specimens 55 cm long suggested an alternative explanation (Wolfson, 1983), that the Gulf of Mexico egg wasaborted before term, and that the whale shark is normally ovoviviparous. This was recently confirmed by an adult femalewhale shark caught in Taiwan (Province of China) which had some 300 young (sex ratio of 237 young with about 1.0:1.1male:female ratio) in her uteri (Joung et al., 1996). These young whale sharks were three size classes: embryos with yolksacs in egg cases that were 42 to 52 cm long, embryos with yolk sacs in egg cases 52 to 58 cm long, and apparent termfoetuses without egg cases and with reabsorbed yolk sacs between 58 and 64 cm long. The type of ovoviviparity practisedby the whale shark is possibly a relatively simple sort very similar to that of the related nurse sharks (Ginglymostomatidae:Ginglymostoma), with retention of the egg case in utero until the embryo hatches from it, and then is born. The three classesof young reported in the Taiwan (Province of China) female suggest that successive batches of eggs are retained in utero,with the oldest hatching and then being born. One additional adult female whale shark from Taiwan (Province of China) wasrecorded as having 16 egg cases in her uteri while another was reported as having 200 eggs in her ovary. Although manywhale sharks have been caught in the Taiwan (Province of China) fishery, very few have been reported as obviously maturefemales over the last half of the twentieth century (Joung et al., 1996). The gestation period is not known, but Castro,Woodley and Brudek (1999) suggest that the whale shark may reproduce every other year as with the nurse shark(Ginglymostoma cirratum).The smallest free-living whale sharks are 55 to 59 cm long, some of which have an umbilical scar. Such small whale sharkshave been found off tropical West Africa in the East-Central Atlantic and near Central America in the eastern Pacific, nearcontinental waters and in the open ocean far from land (Wolfson, 1983; Kukuyev, 1996), suggesting that young may be bornin the ocean and that pupping grounds and possibly nursery areas exist there.The whale shark is a versatile suction filter-feeder, and feeds on a wide variety of planktonic and nektonic organisms. Whalesharks are known to appear off coral reefs when these are producing blooms of planktonic organisms and the corals arespawning (Colman, 1997). Masses of small crustaceans (including copepods) are regularly reported as food, along withsmall and not so small fish such as sardines, anchovies, mackerel, and even small tunas and albacore as well as squid.Whale sharks may aggregate along with tuna in association with spawning of lanternfish (Diaphus, Myctophidae) in the

206 FAO Species Catalogue for Fishery Purposes No. 1Coral Sea off Queensland; and off Christmas Island in the eastern Indian Ocean between Java and Western Australia inassociation with mass spawning of red crabs (Geocarcoidea natalis; Colman, 1997). Recent research (Heyman et. al, inpress) indicates that large (ca. 25 individuals) and predictable groups of whale sharks gather around snapper (Lutjanuscyanopterus and L. jocu) spawning aggregations at Gladden Spit, in the barrier reef off Belize. Whale sharks feed at duskand dawn on the released gametes of the snappers, during the full moon periods between April and June and show goodsite fidelity between one spawning period and the next.The whale shark feeds at or close to the surface, and often assumes a vertical position with its mouth up above its body.Phytoplankton often occurs in the stomachs of whale sharks, but whether this is a major component of the diet of this shark israther doubtful. Small whale sharks 3.2 to 5.2 m long have been observed feeding on copepods at the surface in the Gulf ofCalifornia (Clark and Nelson, 1997); they aimed at patchy areas of dense concentrations of copepods (at least 13 species,mostly Acartia clausi) and turned from side to side, lifted the dorsal surfaces of their heads partially out of the water with theupper jaw exposed, and opened and closed their mouths and gill openings at rates of 7 to 28 times per minute, apparentlygulping in plankters. Turning movements increased with gulping rate, and, when sharks passed through copepodconcentrations into clear water, they sharply turned and circled back to the concentrations to renew feeding. When notfeeding the sharks dropped their heads below the surface with mouth slightly open, and stopped moving their mouths andgill openings and swam faster while apparently ram-ventilating their gills. Larger whale sharks 6 to 10 m were observed byClark and Nelson to feed underwater with their heads not exposed, but with a similar gulping action of 16 to 20 times perminute. Clark and Nelson also noted that similar feeding behaviour is shown by whale sharks kept at the Okinawa ExpoAquarium, fed by ladle at the surface. Some air may be swallowed during feeding, but it is unknown if it is expelled or canfunction for assisting buoyancy as in the sand tiger sharks (Carcharias taurus).The suction-filter mechanism of the whale shark is more versatile than the dynamic filter mechanism of the basking shark inthe range of prey species that can be taken. The basking shark, with its huge scoop-like mouth, hydrodynamically ‘clean’ gillrakers, and huge gill slits, has little if any suction capacity and must depend for the most part on its relatively slow forwardmotion to carry animals into its mouth; this limits it to eating small planktonic crustaceans and other invertebrates trapped onmucus on its gill rakers. The whale shark is not dependent on forward motion to operate its filters, and can probably achieverelatively high intake velocities of water into its mouth, that enable it to readily ingest larger, active nektonic prey in addition tomasses of planktonic crustaceans. A disadvantage of the suction plankton feeding of the whale shark over the dynamicmethod used by the basking shark is that the structures involved can filter a far smaller volume of water per unit time andhence are far less efficient in concentrating diffuse plankters. Hence the whale shark may be more dependent on higherconcentrations of plankters than the basking shark to optimally utilize such food (such as reef blooms or copepodaggregations), but has the option of utilizing much larger nektonic organisms for prey that cannot be caught by the baskingshark or are marginal for that species. Observations by Clark and Nelson (1996) suggest that the whale shark is capable ofhoming in on such concentrations and adjusts its activities to target them.The predators of whale sharks are little-known apart from humans. A newborn specimen was found inside the stomach of ablue shark (Prionace glauca; Kukuyev, 1996), and other large pelagic and coastal carcharhinoid and lamnoid sharks maytake young whale sharks also. Adult whale sharks, with their thick hides and great size, may have few natural predators,killer whalea, (Orcinus) and large white sharks being two possibilities; the giant extinct megatooth shark Carcharodonmegalodon is another likely candidate. The chief predator of whale sharks is humanity.Pauly (in press) used a von Bertalanffy growth curve partially based on data for the basking shark (Cetorhinus maximus)togive a tentative estimate of age and growth for this species. Using a conservative maximum length of 14 m he suggested thatthe whale shark may be especially long-lived, with a tentative longevity of about 100 years, “which strikes one as rather high,but may not be impossible” (Pauly, in press). His growth curve suggests that adult males at about 7 m would be about 20years old. In contrast, Castro, Woodley and Brudek (1999) suggested that, based on captive growth of a term foetus inTaiwan (Province of China) from the adult female reported by Joung et al. (1996), as well as the few records of small whalesharks in the 1 to 3 m range, the whale shark may prove to be the fastest-growing shark. Long-term tagging and bodyprintingof free-ranging individuals with periodic remeasuring of registered individuals that return to viewing sites over several years(as per the methodology of Stevens et al., 1997), may clarify this seeming disparity, and the writer suspects that very fastinitial growth (if also shown by free-living neonates) would slow down markedly as the sharks approach maturity.A preliminary DNA analysis of cytochrome b genes from several whale shark skin samples from the Gulf of California, and afew from Philippines and South Africa, showed no variation (Eckhart and Stewart, 1996). Further studies are underway withother genes to determine if there are any genetic indications of differences within whale sharks from a given area or fromdistant areas.Size: This is by far the world’s largest fish-like vertebrate, with an uncertain maximum size. Old sight records as well asrecent tagging studies and whale shark fishers’ reports suggest a maximum length of 17 to 18 m or even 21.4 m. Specimensare uncommon above 12 m, and 30 specimens reported from South Africa by Beckley et al. (1997) were 4 to 11 m long. Alength of 13.7 m is often given as the maximum size measured, 12.1 m the most recently accurately measured, while mostreported in the literature are between 3 and 12 m long. The late Margaret M. Smith showed the writer a letter describing abeach-stranded specimen from Angola, with measurements suggesting it was about 15.9 m long.Size at birth of the whale shark is between 55 and 64 cm, with freeliving individuals as small as 55 and term foetuses knownat 58 to 64 cm. Males are immature at 299 cm or less and adolescent at 390 to 540 cm while adult males of 705 to 1 026 cm

Sharks of the World, Vol. 2 207have been recorded. Females from 340 cm or less, to 760 cm, were immature, while a pregnant female was about 10.6 mlong (Joung et al. 1996) and weighed 16 t; and another adult female may have been about 12 m long.Pauly (in press) assumed a maximum length of 14 m and a weight of approximately 20 t for the whale shark using anisometric length-weight equation also used with the basking shark: W(t) = 0.0075TL(m) 3 . Recent records suggest a highermaximum length, however. Eckert and Stewart (1996) tracked 12 whale sharks by satellite tags for which size estimateswere given to a tenth of a metre; 10 of their tagged sharks were 3.0 to 7.1 m long, but two big females tagged in 1996 andtracked for four months were 15.0 and 18.0 m long. Taiwanese fishers reported several whale sharks between 15 and 36 tweights (Joung et al., 1996), suggesting lengths of about 12 to 17 m using Pauly’s equation.Interest to Fisheries and Human Impact: Apparently of limited value for conventional fisheries despite expanding marketsand increasing values for whale shark products. Small harpoon fisheries traditionally existed in Pakistan and India for localutilization; it is also taken by harpoon in the Maldives, China, Taiwan (Province of China), and Philippines, and has beencaptured and utilized in Senegal; it is also caught as a bycatch in fish traps in Philippines but was generally released until itincreased in value and was killed for export, and hopefully is now being released again following nation-wide protection.More importantly, an increase in demand for whale shark meat in Taiwan (Province of China) stimulated the development ofa targeted fishery for whale sharks developed by ex-whalers operating in the Bohol Sea. Whaling harpoons or gaffs wereused to subdue them and knives (bolos) to kill them (WWF-Philippine Program, 1996). It is also caught with longlines and ingill nets in Taiwan (Province of China).Whale shark meat is eaten by people fresh, fresh-frozen, dried or dried-salted, the skin is eaten in Taiwan (Province ofChina), the fins enter the oriental fin trade at a high value because of their size, the dried gill rakers have been utilized in thePhilippines, and the flesh has been used to treat boat hulls in Pakistan.The whale shark is generally considered harmless despite its size, and moderate-sized to very large individuals have beenrepeatedly approached closely by divers and have been touched, ridden and otherwise contacted by them without thesharks reacting aggressively. They may suddenly dive or flee the vicinity of divers when disturbed but without showing muchexcitement. Their docility and ready access in shallow water in many localities has popularized them as the subject ofecotouristic diving charters. The best known site is off Ningaloo Reef (Ningaloo Marine Park) in Western Australia, but sitesalso exist off the KwaZulu-Natal coast of South Africa, off Mozambique, Kenya, Seychelles, Thailand, Philippines, theHawaiian Islands, the Gulf of California (Mexico), the Pacific coast of Costa Rica and Colombia, Chile, the Gulf of Mexicocoast of the United States (Texas, Florida), and Belize. These sites allow divers to examine whale sharks underwater, and insome instances (Ningaloo Marine Park) access to whale sharks is restricted by a strict code of conduct limiting interferencewith these sharks by divers and boats (Colman, 1997). The effect of ecotourism on the behaviour and local abundance ofwhale sharks off Western Australia is currently under detailed study (Stevens et al., 1997). The whale shark dive industry ishighly regulated by the Western Australian government, with limited numbers of operators and vessels (16 in 1993decreasing to 14 in 1996 according to Colman, 1997). Operators require commercial tourist licences with yearly or two-yearrenewal, and pay a fee per tourist per day. A benefit of ecotouristic activity with whale sharks, particularly in WesternAustralia but also in the Gulf of California, South Africa and elsewhere, is a rise in scientific activity focusing on the whaleshark, often in cooperation with ecotouristic dive operators.There have been a few cases of whale sharks butting sportsfishing boats (Smith, 1967), possibly after becoming excited byhooked fishes being played from the boats or by bait, but ordinarily they do not contact boats although they may investigatethem very closely. Far more commonly human beings inadvertently ram whale sharks with ships and boats as the sharksbask or swim on the surface (documented in numerous papers by Eugene W. Gudger, cited in Wolfson and de Sciara, 1981).During 1998, a whale shark swam into and became caught in a cooling water intake of a coastal nuclear power station atKoeberg, Western Cape, South Africa. The affected reactor had to be shut down and boats and divers were called in toextract the shark. The shark was still alive after about a day and divers ‘walked’ it until it increased its vigour and swam away!The shark may have been attracted to the warm plume of water released by the power plant, and whale sharks have beensighted in the immediate vicinity of the powerplant before.The whale shark has been kept in captivity in Japan and Taiwan (Province of China) and is relatively hardy if properly fed andhandled. At least 14 largish (3.9 to 6.3 m) individuals have been kept in Japan, primarily in a large oceanarium tank at theOkinawa Expo Aquarium (Clark and Nelson, 1997) for extended periods of over a year. These have learned to feed at thesurface of the tank when presented with a long-handled ladle filled with food such as euphausiid shrimp, squid, and fish, andso fed do not require planktonic food in their tank. More recently term foetuses from a pregnant female whale shark weresuccessfully kept in captivity in Taiwan (Province of China) and Japan.The conservation status of the whale shark is of major concern to scientists and to the public, with expanding fisheries andincreasing value of whale shark products such as flesh and fins in Philippines, India, Taiwan (Province of China) andelsewhere in the late 1990s. This runs counter to the increasing international popularity of live whale sharks as subjects ofecotouristic dives, as well as increasing public sympathy for these animals worldwide as harmless “gentle giants” or “gentlemonsters of the deep” (Clark, 1992) that (as with cetaceans) should be conserved because of their intrinsic worth andemotional appeal.Although whale sharks have been caught off Taiwan (Province of China) for many decades of the twentieth century, whaleshark flesh became very popular fresh for human consumption in Taiwan (Province of China) over the past two decades,which caused a major increase in the value of whale shark products there, encouraged Taiwanese fishers to catch more

208 FAO Species Catalogue for Fishery Purposes No. 1whale sharks, apparently caused a decline in catches of whale sharks off Taiwan (Province of China), but also stimulated thefishing of whale sharks in Philippines until the fishery declined and it was banned in 1998. The Taiwanese market has alsostimulated expanded and substantial export fisheries for whale sharks off India, which developed from traditional artisanalfisheries there and which are apparently declining at present due to overfishing. Whale shark meat was valued at about 400New Taiwanese Dollars per kilo in 1996 (Joung et al., 1996; equivalent to US$13 in 2000 and nearly US$200 000.00 for themeat alone from a 36 t shark if two-fifths of its weight is muscle). Fin prices are uncertain but are probably very high atpresent as with fins of other large sharks.It is uncertain what effects the now banned drift net fisheries had on whale sharks (and for that matter mantas and devil rays)as discarded bycatch during the peak period of their use during the late 1980s and early 1990s. No countries that reportshark catch statistics to FAO currently report whale shark catches.The whale shark has been listed on the IUCN Red List for the past few years, is protected by the United States east coastshark management plan, and in the Maldives. More recently (1998), it was protected by the Philippine government with banson killing and selling them following steep declines in numbers in Philippine waters; similar protection was given to thisspecies off Gladen Spit, Belize (2000). Ecotouristic viewing of whale sharks is being actively promoted by the World WideFund for Nature as an economic alternative to whale shark fisheries in Philippines (M.N.R. Alava and A.A.S.P. Yaptinchay,pers. comm.). After lobbying by the US Shark Research Institute and the Hubbs-Sea World Research Institute the Hondurangovernment declared total protection for the whale shark off Honduras in 1999 (M. Levine, pers. comm.). The species isunder consideration for total protection in South Africa also (1999 to 2000). International protection for the whale shark isprobably necessary, perhaps in the form of a CITES listing with a worldwide ban on fishing them for international trade aswell as regional agreements and national regulations for protecting them or limiting exploitation. Whale shark fins and meatfall into international trade and regulation, but the highly migratory whale shark ranges close inshore and is also subject tolocal fisheries supplying local markets as well as pelagic fisheries and international exporters.There is considerable concern that whale sharks are extremely vulnerable to overexploitation due to their relatively lowabundance, large size, ease of access at the surface, and possibly very slow growth and exceptional longevity (see above).They are a ready target of coastal and pelagic fishing operations, and are easily harvested by small boats in shallow coastalwaters. It is a natural extension, as in Philippines, for former whale-hunters to transfer their activities to whale sharks. Thisparallels whale fisheries that also took basking sharks. With the current high values of whale shark flesh and fins, whalesharks could be targeted in international waters by long-range fishing vessels run like miniature whale factory ships (forexample, a few converted ocean-going stern trawlers of modest displacement set up for processing carcasses andblast-freezing the meat and fins) and using small ‘killer’ boats, harpoon-guns, light helicopters or microlight aeroplanes asspotters, and even remote sensing from satellites to fish these sharks pelagically. Small specialist killer boats withharpoon-guns already exist in the declining basking shark fishery, and could be applied to the whale shark fishery forshore-based operations or serving small factory ships. Fortunately this has not happened to date, and hopefully will neverhappen, as the effect of even a few such vessels or a small fleet could be devastating in short order. It may be possible thatdecreases of whale sharks from shore-based small operations may preclude high-technology pelagic whale shark fishing oneconomic impracticality.Pauly (in press) suggested that because of the whale shark’s slow growth and other life-history parameters any plan fortargeted exploitation of whale sharks as fisheries resources would lead to a quick collapse of numbers, and that evenecotouristic viewing of whale sharks on feeding and mating grounds should be carefully monitored to prevent even indirectmortality which these sharks probably could not accommodate. His prediction apparently was verified while his paper was inpress (1997 to 2000), with the steep decline of the Taiwanese and Philippine whale shark fisheries and possible declines ofwhale sharks in the western Indian Ocean following the developing Indian fishery.It is quite possible, however, that whale sharks are far more valuable in terms of long-term and long-range revenuegenerated from ecotourism than current fisheries, which have the potential to decimate them and remove the source ofrevenue. For Ningaloo Marine Park, Western Australia, revenue from whale shark ecotourism has been growing at 15% peryear and is estimated at being worth A$12.8 million in the year 2000 despite being highly seasonal and of short duration,between March and May of each year (Colman, 1997). If the same sharks visit different viewing sites, as is suggested bylong-range tagging and tracking, and are long-lived, they each may generate far more revenue as a live animal viewedrepeatedly by diving ecotourists over several decades and at several sites rather than that received as a one-off fee fromkilling them (and with low value often received by fishers in Developing countries compared to fin and meat dealers in thedeveloped world). This may be particularly important in places such as the Philippine Islands, where ex-whale shark fishersare becoming involved in whale shark ecotourism (M.N.R. Alava and A.A.S.P. Yaptinchay, pers. comm.) and even a whaleshark museum. Extraordinarily high profits from flesh and fins drive the current fisheries, but their existence is probablyephemeral as whale sharks may be unable to sustain them for very long because of their biological limits.Local Names: Basking shark, East Indian basking shark; Mhor, Chagrin; Tiburón ballena, Pintado; Dominó, Tiburón dama(Mexico); Tubarâo baleia, Whale shark (Azores); Chlarm plawarn (Thailand), Jimbeizame, Yasurizame (Japan); Tofu sa(Taiwan (Province of China)); Isdang tuku, Tuk, Tuku, Tuki Tuki (Philippines).Literature: Smith (1828, 1829, 1849); Müller and Henle (1839); Regan (1908a); Garman (1913); Gudger (1915, 1931,1935, 1940, 1941a,b,c); White (1930, 1937); Herre (1925, 1953); Fowler (1936, 1941, 1967a); Denison (1937); Bigelow andSchroeder (1948); McCann (1954); Baughman (1955); Reid (1957); Garrick (1964); Smith (1967); Iwasaki (1970);Compagno (1973, 1984, 1988, 1990a, b); Bass, D’Aubrey and Kistnasamy (1975c); Johnson (1978); Wolfson and de Sciara

Sharks of the World, Vol. 2 209(1981); Cadenat and Blache (1981); Uchida (1982, 1984); Castro (1983); Wolfson (1983, 1986); Nakaya and Shirai (1984);Dingerkus (1986); Sadowsky et al. (1986); Uchida, Toda and Kamei (1990); Au (1991); Clark (1992); Anderson and Ahmed(1993); Michael (1993); Last and Stevens (1994); Gifford (1994); Seret (1994); Randall (1995); Wamukoya, Mirangi andOttichilo (1995); Joung et al. (1996); Eckert and Stewart (1996); WWF-Philippine Program (1996); Kukuyev (1996); Beckleyet al. (1997); Bonfil (1997); Clark and Nelson (1997); Colman (1997); Santos, Porteiro and Barreiros (1997); Stevens et al.(1997); McEachran and Fechhelm (1998); Castro, Woodley and Brudek (1999); Gunn et al. (1999); Heyman et al. (in press);Pauly (in press); M.N.R. Alava and A.A.S.P. Yaptinchay (pers. comm.); M. Kroese (pers. comm.); M. Levine (pers. comm.);S. Uchida (pers. comm.).There is currently an annotated whale shark bibliography at the Scripps Institution of Oceanography website by Fay HenryWolfson and Giuseppe Notarbartolo-di-Sciara: http://scilib.ucsd.edu/sio/indexes/whalshrk.html.click for next page

click for previous page210 FAO Species Catalogue for Fishery Purposes No. 13. LIST OF SPECIES BY MAJOR FISHING AREASpaGEOGRAPHICAL DISTRIBUTIONMajor Areas for Statistical PurposesSpecies g Fresh 18 21 27 31 34 37 41 47 48 51 57 58 61 67 71 77 81 87 88e water ARC WNA ENA WCA ECA MED WSA ESA ANC WIO EIO ANE WNP ENP WCP ECP WSP ESP ANWAlopias pelagicus 81 Alopias superciliosus 83 Alopias vulpinus 86 Brachaelurus waddi 145 Carcharias taurus 58 Carcharodon carcharias 100 Cetorhinus maximus 91 Chiloscyllium arabicum 167 Chiloscyllium burmensis 168 Chiloscyllium griseum 169 Chiloscyllium hasselti 171 Chiloscyllium indicum 172 Chiloscyllium plagiosum 173 Chiloscyllium punctatum 175 Cirrhoscyllium expolitum 133 Cirrhoscyllium formosanum 134 Cirrhoscyllium japonicum 135 Eucrossorhinus dasypogon 151 Ginglymostoma cirratum 192 Hemiscyllium freycineti 179 Hemiscyllium hallstromi 180 Hemiscyllium ocellatum 181 Hemiscyllium strahani 182 Hemiscyllium trispeculare 183 Heterodontus francisci 36 Heterodontus galeatus 38 Heterodontus japonicus 39 Heterodontus mexicanus 41 Heterodontus portusjacksoni 42 Heterodontus quoyi 45 Heterodontus ramalheira 46 Heterodontus sp. A 49 Heterodontus zebra 48 Heteroscyllium colcloughi 147 Isurus oxyrinchus 109 Isurus paucus 115 Lamna ditropis 119 Lamna nasus 121 Megachasma pelagios 75 Mitsukurina owstoni 69 Nebrius ferrugineus 196 Odontaspis ferox 64 Odontaspis noronhai 66 Orectolobus japonicus 154 Orectolobus maculatus 155 Orectolobus ornatus 158 Orectolobus sp. A 161 Orectolobus wardi 159 Parascyllium collare 137 Parascyllium ferrugineum 138 Parascyllium sp. A 141 Parascyllium variolatum 140 Pseudocarcharias kamoharai 72 Pseudoginglymostomabrevicaudatum 200 Rhincodon typus 203 Stegostoma fasciatum 186 Sutorectus tentaculatus 162

211 Sharks of the World, Vol. 24.1 Keeping Sharks for Scientific Study4. APPENDIX4.1.1 Generalities About Shark CollectionsMany countries have natural history museums, and these often have fish research collections with sharks in them. Thesecollections are essentially ‘libraries’ of preserved specimens, which have a wide variety of uses, specifically in systematic,morphological and phylogenetic studies but more generally for natural history, biology, ecology, feeding, reproduction,parasitology, distribution, conservation, and other disciplines. Although sharks form a moderately large and important groupof fishes, they are poorly known systematically and biologically, and many described species are inadequately representedin museum collections. Also, new species are still being collected at a good rate, especially deepwater benthic speciescollected by trawl and longline gear on continental and insular slopes, seamounts, submarine ridges, and other topographicfeatures in the open ocean, as well as the offshore and inshore tropics of the Indo-West Pacific. It is not impossible thatreaders of this catalogue may come upon unusual sharks including rare species that are interesting and important to scienceor even new species, and may want to save them for later identification, deposition, and study in a museum collection.Hence it is desirable to outline conventional methods of preparing sharks for scientific study along with some background ontheir use.In the eighteen and nineteenth centuries sharks intended for research collections were often skinned and their skins driedand either stuffed or mounted flat (like herbarium specimens) and stored dry along with shark jaws, sawfish saws, stingrayspines, and other dried skeletal parts. As the years went by, dry storage of whole specimens was supplemented and largelyreplaced by wet storage, which is more satisfactory for systematic collections because of less dehydration damage andready accessibility of internal structures through dissection, clearing and staining, radiography, and other techniques.Specimens are prepared first by fixation, stopping putrefaction in fresh material using a powerful antiseptic solution thatmodifies fresh tissues to stable forms that are resistant to agents of decay; and preservation, long-term storage in anantiseptic solution. Ordinary table salt (Sodium chloride, NaCl) was used as brine (high concentrations of salt in water) to fixor ‘pickle’ whole wet specimens or skinned specimens, and also used as a drying and antiseptic agent to pack, dehydrate,and fix dry specimens. High concentrations (70% or more) of ethyl alcohol (CH 3CH 2OH) in water, including high-proofalcoholic drinks, served to fix and preserve whole specimens. Salt and strong alcoholic drinks are old substitutes for fixingand preserving specimens when nothing else is available.Traditional fixation and preservation. By the beginning of the twentieth century fixation in water solutions of formaldehydegas (CH 2O) or formalin, followed by preservation in aqueous solutions of ethyl, isopropyl (CH 3CHOHCH 3) or n-propyl(CH 3CH 2CHOH) alcohols became the standard mode of preparation and storage of whole wet fish specimens includingsharks. The preferred preservative for such collections is 70 to 75% ethyl alcohol, but it has the disadvantages ofexcessively dehydrating some material such as skeletal parts, is highly flammable, and may be more expensive and moresubject to tariffs and storage regulations than other alcohols because of its consumption by people. Isopropyl or n-propylalcohols (particularly the former) in 50% aqueous solution may be more suitable for storage of larger specimens over 60 cmlong or wide, but are less suitable for preserving smaller whole specimens below that size. This particularly applies to smalldelicate specimens because of poorer long-term preservation. Large alcohol-preserved collections may present a firehazard in some countries with histories of seismic activity, volcanism, and large urban fires resulting from natural andhuman-induced disasters. Alcohol collections are costly to maintain due to the expense of alcohol, its high evaporation rate,the need to provide facilities with adequate ventilation and fire protection, and the need for constant vigilance and service toprotect these collections from dehydration. Hence some collections preserve sharks and other fishes in weak formalin,sometimes buffered with hexamine or carbonates. However, formalin preservation is far less desirable than alcohol becauseof the excessive hardening of soft parts including fins, decalcification of hard tissues, and the toxic hazard and nastiness ofworking with formalin. Formalin fixation and preservation in ethyl alcohol is best for long-term maintenance of specimenslarge and small, with isopropyl a substitute for ethyl alcohol when ethyl alcohol is prohibitively priced or regulated.Alternatives for fixation and preservation. A 1% solution of propylene phenoxcetol with 5% ethylene glycol in water hasbeen used in some museums as a substitute preservative for formalin-fixed specimens. In some instances phenoxcetolpreservation has been excellent, but in others specimens began to decay because the phenoxcetol lost its antisepticqualities. Non-aqueous, antioxidant and antiseptic fixing and preserving fluids have been used experimentally for preservingthe life colours of tropical reef fishes, but have not been used much with sharks. There is at least one institution that has triedlong-term preservation of sharks by storing them in freezers, but these specimens became dehydrated and mummified afterlong storage.Dried shark parts may be useful for identification if nothing else is available. Shark parts such as jaws, saws, and stings, aswell as small whole sharks, can be dried in sunlight or indoors at room temperature or with artificial heat sources. Jaws andother shark parts should be thoroughly washed of blood and have most skin and muscle removed before drying. Formalin,alcohol, brine fixation, or salting of the cleaned parts before drying will retard bacterial growth and insect damage, but ifspecimens are to be dried without fixation they should be protected to keep out insects. Alternatively, ant colonies in the fieldand dermestid beetle colonies in museum facilities can be used to remove flesh from semi-dried but unfixed shark jaws orother parts. Likewise marine isopods can be used to remove muscle from fresh jaws or whole skeletons hung in perforatedcontainers from piers in harbours or placed on the bottom in protected waters such as pools. Cleaned shark jaws should bepinned to a board (a large piece of Styrofoam or rubberised foam will suffice) or tied to a frame to keep them from becomingdistorted as they dry.

212 FAO Species Catalogue for Fishery Purposes No. 1Freeze-drying whole sharks may produce better results than stuffing them for display, but is not utilized in systematiccollections to any extent and has several disadvantages over conventional wet-storage of sharks including stiffness andproblems with studying internal structures. Making moulds of specimens can produce high-fidelity casts of whole animals orparts for museum displays and as museum specimens, but as with stuffed specimens these have less utility than wetmaterial.Small formalin-fixed sharks including foetuses are suitable for clearing and staining of skeletons with various techniques.The writer prepares skeletal material from fresh or thawed-frozen sharks of all sizes by maceration of soft tissue in boilingwater and dissecting and washing off the tissue. The cleaned skeletal material is then fixed and preserved in 50% isopropylalcohol to avoid the brittleness of formalin-fixed cartilage. There are several other techniques for preparing skeletal materialincluding wax-impregnation of skeletal parts for dry storage (see Compagno, 1988).Tissue samples for DNA and protein analysis are best obtained from fresh material that is frozen and stored at lowtemperatures or, depending on techniques, fixed in full-strength alcohol (ethyl or isopropyl). Alcohol-fixed and preservedwhole specimens and dried fresh material such as shark jaws or sawfish saws can be used for some biochemicaltechniques, but not formalin-fixed material.4.1.2 Practical Advice for Creating Shark CollectionsCollection and handling of specimens. Most smaller sharks a meter or less long, including small species or smallspecimens of larger species, can be readily fixed and even preserved intact in the field as well as in a dedicated laboratory. Ifit is not possible to fix and preserve whole large specimens, their heads, fins and vertebral columns can be readilyaccommodated in barrels and other containers. To prepare a large non-batoid shark for compact storage measure first itstotal length and other basic measurements (see section 1.2.2 Measurements Used for Sharks). Remove its viscera andmost of its muscle mass from the pectoral fin bases to the second dorsal and anal fins with a knife, leaving a long dorsal stripof skin connecting the head to the first dorsal fin, second dorsal, caudal peduncle and caudal fin. Leave also a short ventralstrip of skin connecting the pelvic-fin bases with the anal fin and caudal peduncle. Strip the vertebral column of excess fleshand cut it off at the head and caudal peduncle, cut it into sections if necessary, tie labels to it and the rest of the shark beforefixing it. Wide and very large batoids are often a problem because of their awkward shape. Their size can be reduced byskinning and removing muscle from the pectoral fins, cutting off the pectoral fins and cutting them into sections, andpreserving the head, body and tail either intact or divided into pieces.For best results sharks should be fixed with formalin as soon after death as possible, though they can be frozen or evencovered with ice to halt or retard putrefaction until the specimen can be fixed or delivered to a museum or other facility forfixation and preservation. Excess freezing will dehydrate unprotected specimens, and sharks that are to be frozen forconsiderable periods should be sealed with some water in plastic bags. In hot climates it is especially important to fix orfreeze specimens quickly, as they can deteriorate in a matter of hours after being caught. Tropical markets are excellentplaces to collect inshore and sometimes offshore and deep-water sharks, as fishers often land most of their catch includingoddities, but market specimens should be processed as soon as possible as they may not be in the most fresh condition aftercapture. Specimens should be kept cool, in the shade, and iced or covered with wet cloth or fibrous sack material if theycannot be immediately fixed or frozen. Fast blast-freezing to -40°C is preferable to slower and higher-temperature freezingmethods where putrefaction may proceed apace during the freezing process. Some deepwater sharks with large livers andhigh squalene content may not properly freeze in higher-temperature low-capacity freezing units. Frozen specimens areideal for skeletal preparations (see Compagno, 1988), but after thawing and fixation as whole specimens they often do notlook quite as good as those that were formalin-preserved in the field while fresh.Containers for fixing and preserving sharks. Containers are necessary to fix sharks in formalin, and include jars, plasticbuckets and boxes with lids, small metal barrels, the ubiquitous and durable plastic industrial barrels with tight-sealing lids,cardboard waterproof drums (‘liquipaks’), elongated rectangular tanks, garbage cans or whatever waterproof containersare available locally. Formalin will quickly corrode ordinary steel containers, so these should be either stainless or shouldhave acid-proof coatings if used for fixation or long-term storage. Ideally a long tank or deep tray, or, for wide animals, along and broad tank of wood, plastic, fibreglass, stainless steel or other formalin-resistant material should be used forpreserving sharks in a straight, spread-out position, but it may be possible only to fix and store specimens in cylindricalcontainers in a curled position. Large, heavy plastic bags have been used to fix sharks quite successfully when othercontainers were not available, and very large specimens could be fixed in a temporary container made from a liner ofheavy plastic sheeting and a box or even a hole in the ground. Containers for fixation and temporary storage should havetight-fitting lids to prevent escape of toxic formalin fumes.Small sharks or their parts can be stored permanently in alcohol in glass bottles in sizes from about 300 ml to 4 or 5 l. Plasticbottles are better for fixation and field storage of small specimens but are not as good as glass for permanent storage. Largerglass bottles and tanks from 10 to 20 l are presently difficult to obtain, but plastic pails or buckets up to about 20 l and smallcommercial barrels over 20 l are the modern substitutes. Lids on all containers with preservatives should be tight-fitting toretard evaporation. Plastic lids for screw-top bottles with full-turn threads work well, as do glass lids with wire bails andrubber gaskets on special preserving jars for fruit and vegetables; screw-top lids will allow evaporation, and metal ones willcorrode, unless provided with a good plastic liner.

Sharks of the World, Vol. 2 213The larger sharks, 1.5 m or more in length or width, present special problems for long-term preservation in alcohol. Theyrequire facilities with large durable rectangular or square tanks of fibreglass, composite wood and fibreglass, polyethylene,or stainless steel. Large preservation tanks are often custom-built, but less expensive industrial tanks for transport orstorage of wet and corrosive materials can be readily substituted for them. Tight-fitting covers for tanks are necessary tominimize evaporation.Tools and materials needed for fixation and preservation. Basic tools and materials that are useful for fixing andpreserving sharks include 40% aqueous (concentrated) formaldehyde, 95% alcohol (ethyl or isopropyl), scalpels, aselection of good knives ranging from small ones with 10 cm blades to large butcher knives with 30 cm blades, a sharpeningsteel, forceps, pointed metal probes, large needles with large eyes for tying tags or suturing up large specimens, 20 to 50 mlhypodermic syringes (preferably plastic and having locking collars to keep the needles from coming off while injectingspecimens), large syringe needles (size 16 or larger), protective clothing including rubber or plastic gloves (including someheavy gloves with long sleeves for ‘tank diving’ to retrieve specimens in large containers), aprons, lab coats, rubber bootsand safety glasses, good quality label paper or card of high rag content that will not tear easily or disintegrate when wet,plastic-impregnated paper that can be written on with pencil, tough tag paper or tie-tags with holes for string, linen, cotton orsynthetic string, cheesecloth, mutton-cloth, or ordinary, non-dyed cloth including old bedsheets and pillowcases,polyethylene plastic bags of various sizes and shapes, pencils, and technical pens with indelible India ink.Recommended fixation and preservation procedures. To fix sharks, prepare a 10% formalin solution by adding 1 partconcentrated formaldehyde to 9 parts of water, in a volume enough to at least cover the sharks in the container selected.Formalin is quite toxic, and should be handled with great care in a well-ventilated place, either outside in the shade or in afacility with a fume hood or a room with powerful extractor fans. Gas masks with formalin-specific filters should be used whenventilation is inadequate.Syringes should be used to inject a quantity of formalin into the body cavity (including the stomach and intestine) and alsothe muscle masses of the body, tail, and fin bases, and the head to preserve the brain. For injection of large numbers of largespecimens pressure garden sprayers or even mortuary suffusion equipment can be adapted for injecting sharks. Dilute 10%formalin can be used for injection, though higher strength formalin, 1:4, 1:2 or even undiluted concentrated formaldehyde isvery effective in preventing putrefaction in hot climates. If a syringe is unavailable make several small holes or slits on sidesof abdomen through to the body cavity of the shark with a knife, scalpel or probe, preferably on the right side (the left side isgenerally used for illustration); even when injecting the shark make at least one small slit on the left side of the abdomen.Position the shark flat on its abdomen with fins spread in the preserving container and add enough dilute formalin to cover it.Fins can be pinned out on pieces of styrofoam or other soft material if necessary. Deepwater sharks and some other speciesmay have large and extremely oily livers, which generally leak oil into fixatives and preservatives. Some plastics, includingcoated label-paper, may have chemical problems with such oil, and it may be desirable to wash out excess oil from the bodycavity before fixation and preservation.With larger sharks in hot climates it may be necessary to use stronger formalin, up to 1:4, for initial fixation, or add moreconcentrated formaldehyde to 10% solution if outgassing from putrefaction is evident. Small sharks below 1.5 m should befixed for at least two weeks to a month, while larger specimens should be fixed for a month or more. Volume of the sharkshould not exceed half of the volume of the preserving fluid. Specimens are then preserved in ethyl or isopropyl alcohol afterfixation, with a brief wash in water to remove excess formalin, and stored in suitable containers.Labels, field numbers and field notes. All specimens should have good-quality labels attached or associated with themthat will survive wetting, movement, transportation, and long-term storage. Make a field label for the shark in pencil onplasticized paper or with a technical pen on high quality, high rag content label paper or card (making sure the ink is drybefore wetting the label. The field label should include a field number, cruise and station numbers if relevant, date ofcollection, locality, collector, species (if known), specimen data including total length, precaudal length, weight, sex andmaturity stage when available. The label should be tied to the shark or placed inside its mouth or inside a gill slit beforefixation. Perforated tag labels with attached string or line should be securely fastened by threading their line through smallholes in the bases of the pectoral fins or other fins. Tags using synthetic string such as nylon, dacron, or polypropyleneshould be tied with secure knots as synthetics are often more slippery than natural fibre.Field numbers of individual collectors serve to link specimens to collectors, expeditions, research vessels, and various typesof data sheets. The writer uses the notation LJVC-YYYY-MM-ddX, using a year/month/day and sometimes letters to makeunique field numbers for individual specimens or batches of specimens caught or collected in the same place and time.These field numbers are added to field locality data sheets or field notebook entries, morphometric and other datasheets,and other material pertinent to the specimens (including photographs), and are kept with specimens that are giveninstitutional catalogue numbers. Data including the species, field accession number, specimen data (including length,weight, maturity and sex), station and cruise numbers if relevant, collection date, collector, locality (including coordinates ifavailable), and habitat data as available (including water temperature, depth, bottom conditions, turbidity, salinity, andoxygen level) should be entered in a field notebook or on a field locality data sheet under the field number.Transport. Specimens can be transported wet over long distances in barrels, liquipaks, cardboard boxes, wooden boxes orother containers if they are properly packed. Each specimen should be wrapped in cloth with a label wrapped with it orattached to it or placed inside the specimen, and the cloth secured with string or elastic bands. It is important to wrap anystructures that can puncture plastic bags, such as fin spines, stingray stings, protruding oral teeth, sawshark and sawfish

214 FAO Species Catalogue for Fishery Purposes No. 1rostra, and extremely rough, thorny specimens with extra cloth or other protective material such as cork or dense plasticfoam. Wrapped specimens are conventionally kept moist by soaking them in formalin or alcohol, bagging them in three ormore layers of plastic bags, and cushioning them with packing material from newspapers and shredded paper to plasticbubble-wrap and Styrofoam packing material. Quite recently (2000) airlines have refused to transport such wet specimensin alcohol, formalin, or even propylene phenoxcetol in ordinary airmail, but now charge exorbitant fees for transport of‘dangerous’ cargo. Specimens may have to be washed of fixatives or preservatives and shipped soaked in water by air,provided airfreight or courier services guarantee fast delivery. Surface freight carriers, including ships, still permit specimensto be transported in fixatives or preservatives.Documentation. If it is not possible to preserve any parts of a shark or even if the animal is preserved intact take black andwhite or, preferably, colour photographs of the entire specimen in lateral view, dorsal and ventral view, and the underside ofits head and pectoral fins, using film cameras with slide or print film, digital still cameras, or video cameras (digital video isparticularly effective). For sharks that will be discarded, remove and dry a strip of teeth or the entire dentition from the upperand lower jaws. At minimum, record the date, locality, depth, collector and any other significant data for the specimen andtake the following measurements with a metre stick or tape measure as indicated in the Plan of the Catalogue: total length(TL), precaudal length (PCR), fork length (FL), preoral length (POR); head length (HDL); eye length (EYL); mouth width(MOW); pectoral-fin anterior margin (P1A); pelvic-fin anterior margin (P2A); first dorsal-fin height (D1H); second dorsal-finheight (D2H); anal-fin height (ANH); and dorsal caudal margin (CDM). With specimens that have expanded pectoral fins,including angel sharks, wobbegongs and batoids, measure the disk width (DWI), the extreme distance across thepectoral-fin apices. Take other measurements if possible, including the full set of measurements for non-batoid sharks listedhere. The writer has additional datasheets with sets of full measurements for various batoids and for chimaeroids, which willeventually be listed as datasheets on the Shark Research Centre web site (http://shark.museums.org.za).The writer is quite willing to help any readers who have shark identification problems, time allowing; and photos,measurements and tooth samples or small whole sharks can be sent to him care of FAO.

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Sharks of the World, Vol. 2 2496. INDEX OF SCIENTIFIC AND VERNACULAR NAMESExplanation of the SystemItalicsItalics: Valid scientific names (genera and species).: Synonyms (genera and species, misidentifications and other combinations).ROMAN : Family names.ROMAN : Names of orders.Roman: Suborders, subfamiles, tribes, and FAO and local names.

250 FAO Species Catalogue for Fishery Purposes No. 1Aacanthias, Squalus .............................5Acanthodian ..................................27Acrodonten ...................................32acronotus, Carcharhinus ........................3acutidens, Negaprion ......................62,198africanum, Poroderma ........................186africanus, Isurus tigris. ........................109Akula gigantakaia ..............................96Akula rogataia ................................33Akula sel devaia ..............................125Akuloobraznye ................................96Akuly domovye ................................68Akuly nastoiaschie .............................96Akuly sel devye ...............................97alatus, Isurus ............................108, 115albimors, Carcharodon. .....................99-100Alecrín .....................................114Alfombrera barbuda ...........................133Alfombrera collareja ...........................137Alfombrera de Taiwan. .........................134Alfombrera japonesa ..........................135Alfombrera jengibre ...........................141Alfombrera mohosa ...........................138Alfombrera tasmánica .........................139Alfombreras .................................130Allopia coda lunga .............................88Allopia volpe marina ............................88Alopeciae ....................................78Alopecias .................................78,80Alopecias barrae ..............................86Alopecias chilensis. ............................86Alopecias longimana ...........................86Alopecias vulpes ..............................86alopecias, Squalus .............................86Alopes ......................................80Alopiadini ....................................78Alopias .............................78-80, 82, 98Alopias caudatus ..............................86Alopias greyi .................................86Alopias macrourus ..........................80,86Alopias pelagicus .......................79,81,83Alopias profundus ..........................83,85Alopias superciliosus ..................79,82-83, 85Alopias vulpes ................................86Alopias vulpinus. ..................79,81-83, 86, 99ALOPIIDAE. ................52,72,75,78-79, 89, 98Alopiids. .....................................53Alopiinae. ....................................98Alopioidea. ...................................51Alopius ......................................80amblyrhynchos, Carcharhinus ..................103amboinensis, Cestracion ........................48American porbeagle ...........................125americanus, Carcharias. ........................58americanus, Odontaspis. ........................58americanus, Squalus ........................58,62Ami........................................107Amia calva ....................................3Anchovy-eater ...............................107Anequim ....................................125Anequin ....................................114Anequin barbatana curta .......................114Anequins ....................................97Angel sharks .........2,15-16, 18, 21, 27-29, 127, 148angiona, Tetraoas .............................91angiova, Tetroras ...........................89-91angustum, Ginglymostoma. .....................190Anomotodon ..................................69ANOMOTODONTIDAE .........................69Antacea ...........................31,51,53,126antarcticus, Mustelus ...........................3Aozame ....................................114appendiculatus, Squalus .......................155Arabian bamboo shark .........................168Arabian carpet shark ..........................167arabicum, Chiloscyllium ...............167-168, 172arenarius, Carcharias .......................58,62Arequim .....................................88argus, Squalus ...............................192Arrequim. ...................................125Asterospondyli .........................31,51,126Atlantic mackerel shark ........................125Atlantic mako shark ...........................114Atlantic porbeagle. ............................125atlanticus, Scoliophis ...........................91atwoodi, Carcharias ..........................100aurata, Hannovera. ............................90Australian cat sharks ..........................131Awozame ...................................114Awozame-zoku. ..............................108BBaglul ......................................188Bakazame ...................................96Bake-aozame ................................117Balavala ....................................173Bambak gorbeh ..............................170Bamboa arábiga ..............................167Bamboa birmana .............................169Bamboa capuchona ...........................182Bamboa elegante .............................172Bamboa estrellada ............................175Bamboa estriada .............................175Bamboa gris .................................170Bamboa hombrera ............................180Bamboa indonesa ............................171Bamboa jaspeada ............................179Bamboa moteada .............................183Bamboa ocelada .............................181Bamboa punteada ............................174

Sharks of the World, Vol. 2 251Bamboas ...................................164Bamboo sharks .................16,24,128, 164-166Banded wobbegong ...........................159barbatus, Chiloscyllium ........................155barbatus, Crossorhinus ....................155, 158barbatus, Orectolobus .....................154-155barbatus, Scyliorhinus .........................155barbatus, Scyllium ............................155barbatus, Squalus. ........................152, 155Barbeled houndsharks ..........................16Barbelthroat carpet shark .......................133Barbelthroat carpet sharks .......................24barrae, Alopecias. .............................86Barred bull-head shark ..........................49Barred shark. .................................49Basking shark . . . 20-21, 52-53, 75, 77, 89-91, 96, 96, 98,103, 204, 206, 208Basking sharks ...........6,52-53, 77, 79, 88, 94, 208Batoid .......................................26Batoid sharks ...............................1,27Batoidea .....................................1Batoids ................1-3, 15, 18, 20, 26-27, 29, 103Bat rays .......................................Be kal mora .................................199Bearded shark ...............................155Bearded wobbegong ..........................152Beaumaris shark .........................121, 125Beinhaakal ...................................96belyaevi, Lamiostoma. .................108, 115, 117bideni, Isurus ................................109Big mouth sharks ..............................78Bigeye ......................................85Bigeye ragged-tooth ............................74Bigeye sand tiger ........................52,66-67Bigeye sand tiger sharks ........................53Bigeye thresher .............................84-85Big-eyed thresher ..............................85Bigeyed thresher shark .........................85birostris, Manta ...............................94Black sand tiger ...............................67Blackbanded bamboo shark. ....................170Blackbanded catshark .........................170Blacktip reef sharks ...........................187blainvillei, Cetorhinus ..........................91Blind shark ..................................145Blind sharks. ..........................24,142-143Blue dog ....................................125Blue nurse ...................................62Blue nurse shark ..............................62Blue pointer .............................107, 114Blue porpoise shark ...........................114Blue shark. ..........................114,125, 206Blue sharks. .................................116Bluegray carpet shark .........................147Blue-grey catshark ............................148Blue-grey shark ..............................148Bluespotted bamboo shark. .....................175Bokee sorrah ................................173Bol’shezhabernye akuly .........................96bonae-spei, Heterodontus ....................42,45Bone shark ...................................96Bonito shark .................................114Bonnethead shark ............................102Bonnetheads ................................103Bottle-nosed shark ............................125Bowfin. .......................................3BRACHAELURIDAE ...........127-128, 131, 142-143Brachaelurus ....................132, 142-146, 190Brachaelurus colcloughi. ...............143, 146-147Brachaelurus modestum .......................145Brachaelurus waddi. ..................143-145, 148BRACHYAELURIDAE .........................142brachyurus, Carcharhinus .......................3Bramble shark ................................29Bramble sharks ...............................27brasiliensis, Isistius ............................77brevicaudatum, Ginglymostoma .........190, 192, 199brevicaudatum, Pseudoginglymostoma .......199-200Brown cat-shark ..............................146Brown shark ..................................62Brownbanded bamboo shark ................175-176Brown-banded bamboo shark ...................176Brownbanded catshark. ........................176Brown-banded catshark ........................176Brownspotted catshark. ........................176brucus, Echinorhinus ..........................29Brugde ......................................96Brugden .....................................96Brygde ......................................96Buffalo hornshark ..............................42Bulkophaaie ..................................33Bull head ....................................40Bull shark .....................................4Bullhead .....................................45Bullhead shark ....................20,33,37,45,50Bullhead sharks ......................1-2, 18, 31-33Bumpytail ragged-tooth shark ....................66burmensis, Chiloscyllium ..................168, 172Burmese bamboo shark ........................169Butanding ...................................188Butterfly ray ..................................16Bych’i akuli ...................................33CCachabeo..................................173Ca cheo beo .................................173Ca mari. ....................................107Ca nham. ...................................173Ca nham nhan ................................62Ca nham nhon ................................62caboverdianus, Ginglymostoma. .................192Cacao lixa. ..................................195

252 FAO Species Catalogue for Fishery Purposes No. 1caerulopunctatum, Chiloscyllium ............174-175Cagnassown de foundo .........................66Cagnea. .....................................96Cagnesca grande .............................107Cagnia .......................66,96,107, 114, 125Calderon. ...................................125California bull-head shark. .......................37californica, Squatina. ..........................37californicus, Heterodontus. ......................36CALLORHINCHIDAE ..........................103calva, Amia ...................................3Can da denti ..................................66Can grossu ..................................107Cane di mare di Messina .......................114Cani di mari .................................114Cani di mer ..................................125Caniscu .................................96,107Cape basking shark .........................90,96capensis, Carcarodon ..........................99capensis, Carcharodon .....................99-100Capidolo .....................................96Carago ......................................96Carcaria feroce. ...............................66Carcarodonte ................................107Carcarodonte di rondelet .......................107Carcarodonte lamia ...........................107CARCHARHINIDAE .............4,21,55-56, 61, 103Carcharhinids .................................56CARCHARHINIFORMES 1-2, 16, 23, 30, 51, 53, 126, 128Carcharhinoid ........................20,24,28,53Carcharhinoid catshark ........................128Carcharhinoid catsharks. .......................128Carcharhinoid galeomorphs .......................1Carcharhinoid sharks ..................17,25,29-30Carcharhinoids. .............22-23, 26-27, 32, 53, 128Carcharhinus .....................21,60,100, 103Carcharhinus acronotus. ........................3Carcharhinus amblyrhynchos ..................103Carcharhinus brachyurus .......................3Carcharhinus hemiodon. ........................4Carcharhinus isodon .........................122Carcharhinus leucas. ...........................4Carcharhinus longimanus ...............4,100, 116Carcharhinus melanopterus. .................4,187Carcharhinus remotus ..........................3Carcharia tauro. ...............................62Carchariae ...................................55Carcharias. ...........51,53,55-57, 59, 62, 71-72, 99Carcharias americanus .........................58Carcharias arenarius. .......................58,62Carcharias atwoodi ...........................100Carcharias cuspidatus ..........................58Carcharias ferox ............................63-64Carcharias griseus. ............................58Carcharias kamoharai .......................57,72Carcharias lamia ..........................99-100Carcharias littoralis. ...........................58Carcharias maso .............................100Carcharias noronhai ...........................66Carcharias owstoni .........................58,62Carcharias platensis ...........................58Carcharias rondeletti. .........................100Carcharias taurus . 3, 6, 15, 52, 56-58, 62, 65-66, 70, 73,97, 99, 103, 113, 198, 206Carcharias tigris .............................109Carcharias tricuspidatus .....................58,62Carcharias verus .............................100Carcharias vorax .............................100Carcharias vulpes ..........................80,86Carcharias yangi ..............................72carcharias, Carcharodon. .............94,97,99-100carcharias, Squalus ........................99-100Carchariicae ..................................51Carchariida ...............................51,126CARCHARIIDAE .........................53,55-56Carchariina ...............................51,126Carcharioidea .................................51Carcharoden. .................................99Carcharoden rondeletii .........................99Carcharodon ...........................3,96-101Carcharodon albimors ......................99-100Carcharodon capensis ......................99-100Carcharodon carcharias ..............94,97,99-100Carcharodon megalodon ....................97,206Carcharodon rondeletii. .....................99-100Carcharodon smithii ..........................100CARCHARODONTIDAE .....................96,98Carcharodontinae. .............................96Carcharoidei ...............................51,53carinatum, Stegostoma. ........................186Carpet shark. .....................24,155, 157, 159Carpet sharks. ....................1-3, 126-128, 149Carpetsharks ................................149Cartilaginous fishes .............................2Cat shark ......40,49,139-140, 146, 170, 175-176, 184Catshark ......................3,146, 170, 173, 175Catsharks ............................16,103, 128Catuloidei ...................................126caudatus, Alopias. .............................86caudatus, Squalus ............................172Cazón ........................................3Cearban .....................................96Centracion ...................................34Centracion zebra ...........................34,48Centracion zebra ..............................39Centraciones ...............................31-32CENTRACIONTIDAE ........................32-33Centracoidei ...............................31-32Centrininae ...................................32CENTROPHORIDAE ...........................28cepedii, Squalus (Lamna) ......................109Cestracion. ................................32-34

Sharks of the World, Vol. 2 253Cestracion amboinensis. ........................48Cestracion francisci. ........................34,36Cestracion galeatus .........................34,38Cestracion heterodontus ........................42Cestracion japonicus ...........................39Cestracion pantherinus ......................34,45Cestracion philippi .........................39,42Cestracion philippi var. japonicus ..............39,48Cestracion quoyi ..............................45Cestracion shark ..............................40Cestracion zebra ...........................39,48CESTRACIONIDAE ............................32Cestracionoidei. ...............................32Cestraciontes ..............................32-33CESTRACIONTIDAE ...........................32Cestraciontini ..............................32-33Cestralion ....................................34cetaceus, Squalus ..............................91Ceteorhinus ..................................90Cethorhinus ..................................90CETORHINIDAE. .........20,52,72,75,79,88-89, 98Cetorhinids ...................................53Cetorhininae ............................88-89, 98Cetorhinoidea .................................51Cetorhinus ..................75,88-90, 98, 103, 128Cetorhinus blainvillei ..........................91Cetorhinus gunneri ............................90Cetorhinus maccoyi .........................89-90Cetorhinus maximus .................77,89-91, 206Cetorhinus maximus norman .....................91Cetorhinus normani .........................89-91Cetorhinus rostratus .........................89-90Cetracion ....................................34Chagrin. ....................................208Chalarm gope. ...............................173Chalarm hin .............................173, 175Chalarm lye .................................175Chalarm seour ...............................175Cheiloscyllium ...........................164, 166Cheloscyllium. ...............................144Cheloscyllium furvum .........................145Chien wên sha k’o .............................68Chienhai chang ...............................79Chienhai chang wei sha k’o ......................79Chilarm seour ................................188chilensis, Alopecias ............................86Chiloscylliinae. ...........................128, 164Chiloscyllium ....128, 145, 164-166, 172-173, 175, 177Chiloscyllium arabicum. ...............167-168, 172Chiloscyllium barbatus ........................155Chiloscyllium burmensis ..................168, 172Chiloscyllium caerulopunctatum .............174-175Chiloscyllium colax ...........................172Chiloscyllium confusum ....................167-168Chiloscyllium dolganovi ....................171-172Chiloscyllium freycineti ........................179Chiloscyllium furvum. .........................145Chiloscyllium fuscum. .........................145Chiloscyllium griseum .........166, 169-170, 172, 175Chiloscyllium hasselti .....................170-172Chiloscyllium indicum ................166, 172, 175Chiloscyllium indicum var. margaritifera ..........174Chiloscyllium indicum var. obscura. ..............171Chiloscyllium indicum var. plagiosa ..............174Chiloscyllium indicum var. plagiosum. ............174Chiloscyllium malaianum ......................179Chiloscyllium margaritiferum ...................174Chiloscyllium modestum. ...................143-145Chiloscyllium obscurum .......................171Chiloscyllium phymatodes ......................172Chiloscyllium plagiosum ..........158, 165, 173, 175Chiloscyllium punctatum. . . 145, 166, 168, 172, 175-176Chiloscyllium russellianum .....................176Chiloscyllium trispeculare .....................183Chiloscyllium tuberculatum .....................166Chiloscyllium tuberculatus .....................172Chimaera ....................................17Chimaeras ...................1-3, 15, 19-21, 27, 103CHIMAERIDAE ..............................103CHIMAERIFORMES .................1-2, 17, 20, 103Chimaeroid ...................................17Chimaeroids ....................1,18,20-21, 23, 28Chimaeroids ..................................1Ching sha k’o ................................202CHLAMYDOSELACHIDAE ....................3,29Chlarm plawarn ..............................208Chondrichthyans ..............................27Chondrichthyes ......................1-3, 17, 20, 28Chrossorhinus ...............................153Chrossorhinus lobatus .........................155Chuich’ih sha k’o ..............................56Chyloscyllium. ...............................166cinerea, Odontaspis ............................58cirratum, Ginglymostoma 122, 190-192, 198, 200-201, 205cirratum, Nebrius. ............................192cirratum, Scyllium ............................192cirratus, Squalus. .........................191-192cirrhatum, Ginglymostoma .....................192cirrhatus, Squalus ............................192CIRRHOSCYLLIIDAE .....................128, 130Cirrhoscyllium .......................130-132, 136Cirrhoscyllium expolitum. ..........132-133, 135-136Cirrhoscyllium formosanum ...............132, 134Cirrhoscyllium japonicum. .............132, 134-135cirrhosum, Scyllium ...........................192Cirriscyllium. ........................132, 143-144cirrosum, Ginglymostoma. ..................191-192cirrosus, Squalus .............................186cirrotum, Ginglymostoma ......................192Cobbler carpet shark ..........................163Cobbler shark ................................163Cobbler wobbegong ...........................163

254 FAO Species Catalogue for Fishery Purposes No. 1COBITIDAE ...................................3colax, Chiloscyllium. ..........................172colax, Hemiscyllium. ..........................172colax, Squalus. ...........................172-173Colclough’s shark .............................148colcloughi, Brachaelurus ...............143, 146-147colcloughi, Heteroscyllium .............143, 146-147Colcloughs shark .............................148collare, Parascyllium .....................137, 139Collared carpet shark ......................137-138Collared carpet sharks .....................130-131Collared catshark .............................138Common Atlantic mackerel shark. ................125Common porbeagle ...........................125Common thresher. .............................88concolor ogilbyi, Nebrodes .....................197concolor, Ginglymostoma ...............191, 196-197concolor, Nebrius .....................191, 195-196Confusing bamboo shark .......................168confusum, Chiloscyllium. ...................167-168Cookiecutter shark .............................77Cookiecutter sharks .........................27,94cornubica, Lamna ............................122cornubicus, Lamia ............................117cornubicus, Squalus ...................117,121-122cornubiensis, Squalus .........................122Cornuda ......................................3Cortez bullhead shark ..........................37Corungun sorrah .........................173, 188Cow-fish .....................................96Cownose ray .................................16Cowshark ................................20,107Cowsharks ....................................2Crested bullhead shark .........................38Crested horn shark. ............................39Crested Port Jackson shark ......................39Crested shark .................................39Crocodile shark ..........................52,72-74Crocodile sharks. ........................21,53,71Crossohrinus ................................150Crossorhinae ................................148CROSSORHINIDAE. ..........................143CROSSORHINOIDAE .........................128Crossorhinus. ........................148, 153-154Crossorhinus barbatus. ....................155, 158Crossorhinus dasypogon. ...................150-151Crossorhinus lobatus ......................154-155Crossorhinus ornatus. .........................158Crossorhinus tentaculatus ......................162cuspidatus, Carcharias .........................58Ddaekayi, Oxyrhina ........................109, 122Dalatias licha ................................58DALATIIDAE. ..............................25,29Damiano ....................................107DASYATIDAE ................................103dasypogon, Crossorhinus ...................150-151dasypogon, Eucrossorhinus ............150-151, 196dasypogon, Orectolobus ....................150-151Death shark .................................107Deepsea bigeye thresher ........................85Deepwater sand tigers ..........................63dekayi, Isuropsis .........................109, 122Demon shark ................................107dentatus, Squalus (Scyliorhinus) .................172Dentudo ....................................114Devil ray .....................................16Devil rays .............................19,77,203devisi, Orectolobus ...........................158Dientuso azul ................................114Dientuso prieto ...............................117Dinotopterini ...........................31,51,126ditropis, Lamna. ................20,97,108, 118-119Dlinnokhvostye akuly ...........................79Dlinnozubye akuly .............................56dofleini, Scapanorhnchus. .......................69Dogfish ....................................3,15Dogfish shark .................................28Dogfish sharks ...........................2,18,28doldi, Nebrius. ...............................197dolganovi, Chiloscyllium ...................171-172Dominó .....................................208Dormilón acebrado .............................48Dormilón boquigrande ..........................47Dormilón búfalo ...............................41Dormilón carenado .............................38Dormilón cornudo ..............................36Dormilón de Galápagos .........................45Dormilón de Omán .............................50Dormilón japonés ..............................39Dormilón toro .................................43Dormilones ...................................33Dundanee. ...................................62Dusky dogfish. ...............................146EEagle ray ....................................16Eagle rays ..................................103East Indian basking shark ......................208ecarinata, Lamna ..............................58Echinorhinus brucus ..........................29El marrago ..................................125Elasmobranchii .........................1-2, 19, 28Elasmobranchs. ...........................1,4,27Electric ray ...................................16Elefante .....................................96Elephant de mer ...............................96Elephant fish. ...............................1,17Elephant shark ................................96Elephantfish .................................103elephas, Selache. ..............................91

Sharks of the World, Vol. 2 255elephas, Squalus. ..............................91Elfin shark. ...................................71Elphin shark ..................................71Emissole. .....................................3Endormi ................................199, 201Eostegostoma ...............................190Epaulette shark ...........................181-182Epaulette sharks. ..........................24,177ETMOPTERIDAE ..............................25Ettee .......................................173Eucrossorhinus ........................3,149-152Eucrossorhinus dasypogon .............150-151, 196Eucrossorhinus ogilbyi .....................150-151EUGOMPHODIDAE ............................55Eugomphodus ...........................55-56, 58Eugomphodus griseus ..........................58Eugomphodus littoralis .........................58Eugomphodus taurus ......................3,58,62Euprotomicrus ................................73Euselachii .....................2,19,23,31,51,126Exoles. .....................................117expolitum, Cirrhoscyllium ..........132-133, 135-136FFalse catsharks .............................3,16fasciatum, Stegostoma .....................185-186fasciatus, Squalus .........................185-186ferox, Carcharias ...........................63-64ferox, Odontaspis .....................56,63-64, 67ferox, Squalus ..............................63-64ferrugineum, Ginglymostoma ................196-197ferrugineum, Parascyllium .................138-139ferrugineum, Scyllium .........................196ferrugineus, Nebrius ...............24,190, 196, 201Fierce shark ..................................66Finbacked catsharks ...........................16Flat shark ....................................16Flat sharks ....................................1Foolish shark .................................96formosanum, Cirrhoscyllium ...............132, 134Fox shark .................................83,88Fox sharks ...................................79francisci, Cestracion ........................34,36francisci, Gyropleurodus ........................36francisci, Heterodontus .............33,36-37, 44, 46Freckled carpet shark. .........................179Freshwater dogfish. .............................3Freshwater stingrays ............................1freycineti, Chiloscyllium .......................179freycineti, Hemiscyllium ...............177, 179, 196freycineti, Scyllium. ...........................179Frilled shark ..................................20Frilled sharks ............................2,20,29Fringe shark .................................155Frog shark ..................................173fulvum, Ginglymostoma ........................192furvum, Cheloscyllium .........................145furvum, Chiloscyllium .........................145fuscum, Chiloscyllium .........................145Fynstert-sambokhaai ...........................88GGab doll ....................................107Galapagos bullhead shark .......................45Galapagos bull-head shark ......................46Galapagos horn shark ..........................46Galea ...................................51,126galeatus, Cestacion ............................34galeatus, Cestracion ...........................38galeatus, Gyropleurodus ........................38galeatus, Heterodontus ......................33,38galeatus, Molochophrys. ........................38Galei ....................................51,126GALEIFORMES ...........................51,126Galeoidea .............................51,53,126Galeoidei ................................51,126Galeomorph sharks ..........................2,26Galeomorpha .......................31,51,53,126Galeomorphi. ..................................2Galeomorphii .................................20Galeomorphs .................................28Galeorhinus. .............................28,103Galeorhinus galeus ..........................6,28Galeus vulpecula ..............................86galeus, Galeorhinus ............................6gangeticus, Glyphis .............................4Ganumu sorrah. ..............................114Gata ....................................42,195Gata atlantica ................................195Gata nodriza. ................................193Gata nodriza atezada ..........................197Gata nodriza rabicorta .........................200Gatas nodriza ................................188Gatas nodrizas ...............................126Gato ......................................3,46Gench. .....................................107Ghost shark ..................................17Ghost sharks .............................1,3,27Giant basking shark ............................96Giant guitarfish ...............................103Giant sleepy shark ............................199Gigantskie akuly ...............................96Ginger carpet shark ...........................141Ginglimostoma. ..............................192Ginglymostoma ....126-129, 185, 188-191-193, 202, 205Ginglymostoma angustum ......................190Ginglymostoma brevicaudatum ..........190, 192, 199Ginglymostoma caboverdianus ..................192Ginglymostoma cirratum 122, 190-192, 198, 200-201, 205Ginglymostoma cirrhatum ......................192Ginglymostoma cirrosum ...................191-192Ginglymostoma cirrotum .......................192

256 FAO Species Catalogue for Fishery Purposes No. 1Ginglymostoma concolor ...............191, 196-197Ginglymostoma ferrugineum ................196-197Ginglymostoma fulvum ........................192Ginglymostoma muelleri .......................196Ginglymostoma rueppelli. ......................196GINGLYMOSTOMATIDAE . 24, 127-129, 157, 187-188, 190,205Ginglymostomatinae. ..........................190GINGLYMOSTOMATOIDAE ................157, 188Ginglymostomatoidei ..........................126GINGLYSTOMIDAE .......................128, 188Gingylostoma ................................192glauca, Oxyrhina .........................108-109glauca, Prionace ..................99,116,121, 206glaucus, Isuropsis ............................109glaucus, Squalus ..........................99,121Glyphis gangeticus .............................4Glyphis spp.. ..................................4Goblin shark ...............................52,69Goblin sharks ........................21,52-53, 68gomphodon, Oxyrhina .....................108-109Grand requin blanc. ...........................100Gray carpet shark. ............................176Gray nurse sharks .............................56Gray sharks ..................................56Great white death .............................107Great white shark .........................100, 107Great white sharks .............................97Grey bamboo shark ...........................170Grey nurse ...................................62Grey nurse shark ..........................3,6,62Grey nurse sharks .............................56Grey reef sharks. .............................103Grey shark ...................................62Grey sharks ...............................21,60greyi, Alopias .................................86Grey-nurse ...................................62Griset ........................................3griseum, Chiloscyllium ........166, 169-170, 172, 175griseum, Hemiscyllium. ........................169griseum, Scylia. ..............................171griseum, Scyllium .........................170-171griseus, Carcharias ............................58griseus, Eugomphodus ..........................58griseus, Hexanchus ...........................101griseus, Odontaspis ............................58gronovianus, Squalus. .........................172Grootoog-sambokhaai ..........................85Grootoog-skeurtandhaai. ........................74Ground sharks ..............................2,30guentheri, Isurus .............................109guentheri, Lamna .............................109Guitarfish ................................16,103Guitarfishes ...................................3Gulf wobbegong ..............................159Gulper sharks .................................27Gummy shark ..................................3Gunneri. .....................................90gunneri, Cetorhinus ............................90Gunneri, Squalus (Cetorhinus) ...................91gunnerianus, Squalus. ..........................91güntheri, Lamna. .............................109Gurry shark. ..................................96Gynglimostoma ..............................191Gyropleurodus .............................33-34Gyropleurodus francisci ........................36Gyropleurodus galeatus. ........................38Gyropleurodus japonicus .......................39Gyropleurodus peruanus ........................45Gyropleurodus ramalheira. ......................46HHaa skieding. ................................107Haabranden .................................125Haamar. ....................................125Haar moer ...................................96Hachiware ...................................85Halaelurus. ..................................22halei, Orectolobus ornatus ..................158-159hallstromi, Hemiscyllium ..................177, 180HALSYDRIDAE. ............................88-89Halsydrus .................................88-90Halsydrus maximus. .........................90-91Halsydrus pontoppidani ......................90-91Hammerhead ..................................3Hammerhead sharks ...........................27Hammerheads .........................16,56,103Hannovera ...................................90Hannovera aurata .............................90Hannoveria ..................................90Hare hongi ..................................107Haringhaai ..................................125hasselti, Chiloscyllium. ....................170-172hemiodon, Carcharhinus ........................4Hémiscylle tacheté ............................140Hemiscylliid orectoloboid .......................128HEMISCYLLIIDAE ....................127-128, 164Hemiscylliinae ...........................128, 164Hemiscyllium ................164-165, 177, 183, 196Hemiscyllium colax ...........................172Hemiscyllium freycineti ...............177, 179, 196Hemiscyllium griseum .........................169Hemiscyllium hallstromi. ..................177, 180Hemiscyllium indicum .........................172Hemiscyllium malayanum ......................179Hemiscyllium modestum .......................144Hemiscyllium ocellatum ...........177, 181, 183-184Hemiscyllium oculatum ........................181Hemiscyllium plagiosum .......................174Hemiscyllium punctatum .......................175Hemiscyllium strahani ....................177, 182Hemiscyllium trispeculare .............143, 177, 183

Sharks of the World, Vol. 2 257Hemiscyllium variolatum. ..................136, 140heptagonum, Scyllium .........................186Heptranchias ..................................4Heptranchias perlo .............................4Herbst’s nurse shark ...........................66herbsti, Odontaspis .........................64,66Herbsts nurse shark ............................66Heringshai ...................................97Herring shark ................................125Heterodontea ..............................31-32Heterodontes .................................34Heterodonti ................................31-32Heterodonticae ................................31Heterodontida. ................................31HETERODONTIDAE ........................20,33HETERODONTIFORMES ...........1-2, 18, 20, 31-32Heterodontina ..............................31-33Heterodontoid. ................................20Heterodontoid sharks ........................18,27HETERODONTOIDAE ..........................33Heterodontoidea ........................31-32, 126Heterodontoidei ...............................31Heterodontoids ...................26,28,32,53,128Heterodontus ...........................33-34, 38Heterodontus bonae-spei .....................42,45Heterodontus californicus .......................36Heterodontus francisci .............33,36-37, 44, 46Heterodontus galeatus ......................33,38Heterodontus japonicus ..................39,48,50Heterodontus mexicanus ..................37,41-42Heterodontus oligostictus .......................42Heterodontus peruanus .........................46Heterodontus portusjacksoni .................42,45Heterodontus quoyi ......................33,45-46Heterodontus ramalheira .......................46Heterodontus seftoni ...........................42Heterodontus sp. ..............................37Heterodontus sp. A ............................49Heterodontus zebra .........................33,48heterodontus, Cestracion ........................42HETERONDONTIDAE ..........................33Heteroscyllium ...................142-143, 146-147Heteroscyllium colcloughi. .............143, 146-147Heulgi .......................................96Hexanchida ..................................31HEXANCHIDAE ................................4HEXANCHIFORMES .....................1-2, 4, 20Hexanchoid ..................................27Hexanchoid sharks. ............................22Hexanchoids. ..............................17,53Hexanchus griseus ............................101Higezame ...............................134, 136Hitokiuzame .................................107Hoe mother. ..................................96Hohojirozame ................................107Holocephalans ................................19Holocephali ......................1-2, 17, 19-20, 28Homer. ......................................96Homianus ....................................90homianus, Squalus .............................91Hooded carpet shark ..........................182Horn shark ................................20,36Horn sharks ..................................33Horned shark .................................37Houndsharks .............................16,103huidobrii, Lamna .............................109Husha k’o ....................................33Huta tax-xmara. ..............................107Hypotremata. ..................................1IIkan tjutjot ...................................170Ikan tjutjot kumbang ...........................173Ikan tjutjot matjan .........................175, 188Ikan tjutjot pisang .............................170Ikan tjutjot tekeh ..............................173Imbestinu ................................96,107Indian sand tiger. ..............................62indicum var. margaritifera, Chiloscyllium. .........174indicum var. obscura, Chiloscyllium ..............171indicum var. plagiosa, Chiloscyllium. .............174indicum var. plagiosum, Chiloscyllium ............174indicum, Chiloscyllium ................166, 172, 175indicum, Hemiscyllium. ........................172indicus, Squalus ......................166, 172-173Indonesian bamboo shark ......................171Indonesian speckled carpet shark ................179Indo-Pacific tawny nurse shark ..................196interruptum, Scyllium plagiosum var. .............174Inuzame ....................................176Isdang tuku. .................................208Isistius ................................73,77,94Isistius brasiliensis ............................77isodon, Carcharhinus .........................122isodus, Squalus. ...............................91Isurida. ..................................51,126ISURIDAE ...........................79,89,96-98ISURIFORMES ............................51,53Isurina. ......................................96Isurini .......................................97Isuro muso acuto .............................125Isuroidea. ....................................51Isuroidei ..............................51,53,126Isuropsis. ...................................108Isuropsis dekayi ..........................109, 122Isuropsis glaucus .............................109Isurus .................96-98, 101, 108-109, 117-118Isurus alatus ............................108, 115Isurus bideni ................................109Isurus glaucus ...............................109Isurus guentheri ..............................109Isurus mako .................................109

258 FAO Species Catalogue for Fishery Purposes No. 1Isurus oxyrhynchus ...........................109Isurus oxyrinchus ......97,103, 108-109, 114, 116-118Isurus paucus. ............108, 110, 114-115, 117-118Isurus spallanzanii. ...........................109Isurus tigris africanus .........................109Itar shark ...................................182Jjacksoni, Squalus ..............................42jacksoni, Squalus portus. .................32,34,42jacksonii. Squalus .............................42Japanese bearded shark .......................155Japanese bullhead shark ........................39Japanese bull-head shark .......................40Japanese carpet shark .........................155Japanese goblin shark ..........................71Japanese horn shark ...........................40Japanese mackerel shark ......................121Japanese ragged tooth shark. ....................74Japanese wobbegong .........................154Japanse bulkophaai ............................40japonicum, Cirrhoscyllium .............132, 134-135japonicus, Cestracion ..........................39japonicus, Cestracion philippi var. ................39japonicus, Gyropleurodus .......................39japonicus, Heterodontus ..................39,48,50japonicus, Orectolobus ........................154japonicus. Cestracion philippi var. ................48Jaquetón blanco ..............................100Jaquetón de ley ..............................107Jaquetones. ..................................96Jimbeizame .................................208Jimbeizame-ka ...............................202jordani, Scapanorhynchus .......................69Jumping shark ...............................107KKabouterhaai .................................71Kalouyon ...................................199Kamohara’s sand shark .........................74kamoharai, Carcharias ......................57,72kamoharai, Odontaspis .........................72kamoharai, Pseudocarcharias. ................71-72Karakusa-ôs .................................159Karikkan schura ..............................173Kau kang sha ................................173Kelb Il-bahar .................................107Kelb-il-bahar abjad ............................107Kersch .....................................107Killer whales ..................................94Kirinotobuka .................................155Kitefin sharks ..............................25,27Kitovye akuly ................................202Knopstert-skeurtandhaai ........................66Koesterhaai ..................................96Komrasi ....................................188Kongarasi ...................................188Koumoune ..................................199Kovrovye akuly ...............................149Kumohada-oose ..............................157Kumohada-ôse ...............................157Kurakakezame ...............................136Kurakake-zame ..............................136LLa faux ......................................88lambarda, qualus (Scyliorhinus) .................166Lamea .....................................107Lameo .....................................107Lamia. ...........................66,107, 117-118Lamia (Carcharodon carcharias) ................118Lamia cornubicus. ............................117Lamia oxyrhincus. ............................109lamia, Carcharias ..........................99-100lamia, Squalus (Carcharhinus) ..................100Lamie ...............................97,107, 125Lamio .......................................62Lamiostoma ...........................96,98,108Lamiostoma belyaevi ..................108, 115, 117LAMIOSTOMATIDAE ........................96,98Lamna. .............88,96-98, 101, 108-109, 117-118Lamna cornubica .............................122Lamna ditropis .................20,97,108, 118-119Lamna ecarinata ..............................58Lamna guentheri .............................109Lamna güntheri ..............................109Lamna huidobrii .............................109Lamna latro .................................109Lamna nasus. ................108-109, 118, 121-122Lamna oxyrhina ..........................108-109Lamna pennanti ..............................122Lamna philippi ...........................118,122Lamna punctata ..........................109, 122Lamna smeriglio ..............................125Lamna whitleyi. ..........................118,122Lamnae ..................................51,53Lamnicae ................................51,126Lamnida .................................51,126LAMNIDAE . 21, 27, 52, 69, 72, 75, 79, 88-89, 96-98, 103, 201Lamnids .....................................53LAMNIFORMES ...........1-2, 21, 51, 53, 72, 75, 126Lamnina .................................51,126Lamninae ....................................98Lamnini ...................................96-97Lamnoid ............................20,24,26,53Lamnoid sharks ............................53,75LAMNOIDAE .................................88Lamnoidea ............................51,96,126Lamnoidei. ............................51,53,126Lamnoids. .................15,21,32,52-53, 72, 128Lantern sharks .............................25,27Laoshak’o...................................96latro, Lamna. ................................109

Sharks of the World, Vol. 2 259Le Carcharodonte lamie ........................107Le grand requin ..............................107Le lamie long nez .............................125Le pélerin ....................................96Le squale a fanons .............................96Le squale nez ................................125Le très grand .................................96Leopard shark ...............................188leucas, Carcharhinus ...........................4lewisi, Scapanorhynchus ........................68Liabhán chor gréine ............................96Liabhán mór ..................................96licha, Dalatias ................................58Linnaean shark. ...............................32Lip sharks ...................................166Lisitska morskayia .............................88Little shark ...................................49littoralis, Carcharias ...........................58littoralis, Eugomphodus. ........................58littoralis, Odontaspis ...........................58littoralis, Squalus ..............................58lixa, Squalus. .................................58Lluynog mor ..................................88Loaches ......................................3lobatus, Chrossorhinus ........................155lobatus, Crossorhinus ......................154-155lobatus, Squalus ..........................153, 155Longfin mako .......................52,97,115-116Longfinned mako shark ........................117longicaudatus, Squalus ........................186longicaudatus, Stegostoma .....................185longimana, Alopecias. ..........................86longimanus, Carcharhinus ...............4,100, 116Long-nosed shark. .............................96Longtailed carpet sharks .......................164Long-tailed shark ..............................88Loutre de mer ................................125Ludia. ......................................125Ludia marraco. ...............................114Mmaccoyi, Cetorhinus .........................89-90maccoyi, Tetroras. .............................91macer, Polyprosopus ...........................91Mackerel porbeagle ...........................114Mackerel shark ....................21,114,121, 125Mackerel sharks . . 1-2, 21, 27, 51-53, 79, 96-97, 108, 118macmillani, Parmaturus .........................4macrodous, Squalus ............................58macrourus, Alopias .........................80,86macrurus, Nebrodes. ..........................196maculatus, Orectolobus ...................155, 163maculatus, Squalus .......................152, 155Madame X ..................................199Mako. ......................................114Mako shark ..................................114Mako sharks .................................108mako, Isurus. ................................109Makos .........................21,52-53, 108, 118Mako-sharks. .................................97malaianum, Chiloscyllium ......................179malaianum, Scyllium ..........................196malaisianum, Scyllium .........................179malayanum, Hemiscyllium. .....................179Mambone ...................................114Mandelhai. ...................................96Maneater shark ..............................107Man-eater shark ..............................107Man-eater sharks ..............................97Man-eaters ...................................97Man-eating sharks .............................97Mangia alice .................................107Mango-tuatini ................................107Manta .................................16,29,94Manta birostris. ...............................94Ma-onaga ....................................88Maou urh sha .................................49maou, Squalus (Carcharias) ....................100margaritifera, Chiloscyllium inducum var. .........174margaritiferum, Chiloscyllium. ..................174marina, Vulpecula ..........................80,86Marracho ...............................114,125Marraco ................................107, 125Marrajo .................................114,125Marrajo carite ................................115Marrajo criollo ............................110,114Marrajo dientuso. .............................110Marrajo negro ................................117Marrajo salmón. ..............................119Marrajo sardinero .............................122Marrajos .....................................96Marraquet ...............................114,125Marrraix ....................................125Marteau ......................................3maso, Carcharias. ............................100Mastinu feru .................................107Mauisha....................................49maxima, Selache ..............................91maximum, Selache .............................91maximus normani, Cetorhinus ....................91maximus, Cetorhinus. ................77,89-91, 206maximus, Halsydrus .........................90-91maximus, Selache. .............................91maximus, Squalus ...........................90-91Meanto .....................................114Mediterranean mackerel shark ...................114Megachasma ..................51-52, 74-75, 89, 128Megachasma pelagios. .........................75MEGACHASMIDAE ...................72,74-76, 89megalodon, Carcharodon ...................97,206Megamouth shark ........................21,75-77Megamouth sharks .................29,52-53, 74, 78

260 FAO Species Catalogue for Fishery Purposes No. 1Megatooth shark. ..........................97,206melanopterus, Carcharhinus .................4,187Melantoun. ..................................125Menchenhai .................................107Menschen fresser. ............................107Merviel fras. .................................107Mesorhipidion sharks ...........................28Mexican horn shark ............................42Mexican hornshark .............................41mexicanus, Heterodontus .................37,41-42Mhor .......................................208Micristodus .................................203Micristodus punctatus .........................203Milandre ......................................3Mitsukuri’s shark. ..............................71mitsukurii, Scapanorhynchus. ....................69Mitsukurina ............................52,68-69Mitsukurina owstoni ........................68-69MITSUKURINIDAE .........................68,70Mitsukurinids .................................53Mitsukurizame ................................71Mitsukurizame-ka ..............................68Mizuwani ....................................74Mizu-wani ....................................74Mizuwani ka ..................................56MOBULIDAE ..............................19,77modestum, Brachaelurus .......................145modestum, Chiloscyllium ...................143-145modestum, Hemiscyllium .......................144modestum, Squalus. ...........................146Mokazame ..................................121Molochophrys ..............................33-34Molochophrys galeatus .........................38monensis, Squalus. ........................121-122Monkey-mouth ...............................188Monkey-mouthed shark ........................188Morgi mawr. .................................125Moro .......................................114Morozame. ..................................114Mosambiekse bulkophaai. .......................48Mozambique bullhead shark .....................48Mudshark ...................................107muelleri, Ginglymostoma. ......................196multimaculatum, Parascyllium. ..........136, 138-139Mustelus .................................3,103Mustelus antarcticus. ...........................3Mustelus spp. .................................3MYLIOBATIDAE ..............................103Myliobatis ..................................103NNadebuka ....................................88Nas llarg ....................................125nasus, Lamna ................108-109, 118, 121-122nasus, Squalus ...........................117,121Nasuta ......................................71nasutus, Odontaspis ............................69naucum, Stegostoma tigrinum ...................186Nebria .....................................195Nebriinae ...................................188Nebris. .....................................195Nebrius. .....126-129, 185, 188-190, 195-197, 199, 202Nebrius cirratum .............................192Nebrius concolor .....................191, 195-196Nebrius doldi ................................197Nebrius ferrugineus ...............24,190, 196, 201Nebrodes ...................................195Nebrodes concolor ogilbyi. .....................197Nebrodes macrurus ...........................196Necklace carpet shark .........................140Negaprion acutidens .......................62,198Nekozame ...................................40Nekozame-ka .................................33Neoparascyllium. .........................136, 139Neoselachian .................................23Neoselachians ........................21-23, 26-27Neoselachii. ..........................1,19,23,27Neoselachii ...................................2Nequim .....................................125Neushaai ...................................125Nez........................................125Nezumezame .................................88Nezumizame ................................121Nezumizame-ka ...............................97Nga man ing-myong ...........................173Nitara .......................................83Niuhi .......................................107normani, Cetorhinus .........................89-91normani, Cetorhinus maximus ....................91noronhai, Carcharias. ..........................66noronhai, Odontaspis ...............55-56, 63-68, 73North Australian wobbegong ....................160Northern wobbegong ..........................160Notorynchus .................................60Nurse shark. .............173, 188, 193, 195, 200-201Nurse sharks . . 24, 128, 157, 187-189, 194-195, 200, 205Oobesus, Triaenodon ...........................198obscurum, Chiloscyllium .......................171Oceanic sand tiger .............................67ocellatum, Hemiscyllium ...........177, 181, 183-184ocellatum, Scyllium ...........................181ocellatus, Squalus ........................177, 181oculatum, Hemiscyllium. .......................181oculatus, Squalus .............................181Odontaspe feroce. .............................66Odontaspe tauro. ..............................62ODONTASPIDAE ..............................56Odontaspide taureau ...........................62Odontaspides .................................55Odontaspidida .............................51,53

Sharks of the World, Vol. 2 261ODONTASPIDIDAE . . 3, 52-53, 55-56, 71-72, 75, 98, 103Odontaspidids .............................53,55Odontaspidoidea ..............................51Odontaspis ....................53,55-58, 63, 71, 75Odontaspis americanus .........................58Odontaspis cinerea ............................58Odontaspis ferox. ...............53,56,63-64, 66-67Odontaspis griseus. ............................58Odontaspis herbsti ..........................64,66Odontaspis kamoharai. .........................72Odontaspis littoralis ...........................58Odontaspis nasutus ............................69Odontaspis noronhai ...............55-56, 63-68, 73Odontaspis owstoni ............................64Odontaspis platensis ........................58,62Odontaspis taurus ..........................58,62Odontaspis taurus ..............................3Odontaspis tricuspidatus ........................58Odontaspis tricuspitatus ........................58Odontaspoidea ................................51Ogilby’s wobbegong ...........................152ogilbyi, Eucrossorhinus ........................151ogilbyi, Nebrodes concolor .....................197ogilbyi, Orectolobus .......................150-151Oil shark ......................................6Oilfish .......................................96Old woman shark ..............................96oligostictus, Heterodontus .......................42Oman bullhead shark ...........................50Onagazame ...............................83,88Onagazame-ka. ...............................79Oorookoolti sorrah ............................188Oose. ......................................155orca, Orcinus. ................................94Orcinus ....................................206Orcinus orca .................................94Orectolobicae ................................126ORECTOLOBIDAE. 18, 126-128, 130-131, 143, 148, 164,184, 188, 196, 201ORECTOLOBIFORMES ......1-2, 16, 24, 126, 128, 131Orectoloboid ................3,15,20,24,26,53,128Orectoloboid sharks ...........................128Orectoloboid whale shark. .......................53Orectoloboidea ...........................126, 128Orectoloboidei ...............................126Orectoloboids . . . 17, 19, 21, 25-28, 32, 53, 127-129, 131,196, 202Orectolobus ........3,60,128, 148-150, 152, 158, 162Orectolobus barbatus. .........................154Orectolobus barbutus. .........................155Orectolobus dasypogon ....................150-151Orectolobus devisi ............................158Orectolobus japonicus ........................154Orectolobus maculatus ....................155, 163Orectolobus ogilbyi .......................150-151Orectolobus ornatus ..................153, 158, 163Orectolobus ornatus halei ..................158-159Orectolobus ornatus ornatus ....................159Orectolobus sp. A. ................149, 153, 161-162Orectolobus tentaculatus .......................162Orectolobus wardi ........................156, 159Ornate wobbegong. ...........................158ornatum, Scyllium ........................158, 174ornatus halei, Orectolobus ..................158-159ornatus ornatus, Orectolobus ...................159ornatus, Crossorhinus .........................158ornatus, Orectolobus. .................153, 158, 163ornatus, Orectolobus ornatus ...................159Ôse........................................155Oshirozame .................................107Ossirina dello spallanzani. ......................114Ô-tenjikuzame ...............................199Ôwanizame ..................................66Owston’s sand shark ...........................62owstoni, Carcharias. ........................58,62owstoni, Mitsukurina. .......................68-69owstoni, Odontaspis. ...........................64owstoni, Scapanorhynchus. ......................69Oxyrhina ................................98,108Oxyrhina daekayi. ........................109, 122Oxyrhina glauca ..........................108-109Oxyrhina gomphodon ......................108-109oxyrhina, Lamna ..........................108-109oxyrhincus, Lamia ............................109oxyrhynchus, Isurus ...........................109oxyrinchus, Isurus. .....97,103, 108-109, 114, 116-118Oxyrrhina. ..................................108Oyster crusher ................................45Oyster-crusher ................................45PPacific angelshark .............................37Pacific bonito shark ...........................114Pacific mako .................................114Pacific mako shark ............................114Panchusha.................................175Panray ......................................16pantherinum, Poroderma. ......................186pantherinum, Scyllium .........................186pantherinus, Cestracion. .....................34,45pantherinus, Squalus ..........................186Papa isengezi. ...............................201Papuan epaulette shark ........................180Parascylliid orectoloboids. ....................17,23PARASCYLLIIDAE. ......15,19,26,127-128, 130, 196Parascylliinae ............................128, 130Parascyllium .............127, 130-131, 136, 138-139Parascyllium collare ......................137, 139Parascyllium ferrugineum. .................138-139Parascyllium multimaculatum ...........136, 138-139Parascyllium sp. A ...........................141Parascyllium variolatum. ..................131, 140Parascylloidei ................................126

262 FAO Species Catalogue for Fishery Purposes No. 1Parmaturus macmillani .........................4Pas sabljas ...................................88Patings ......................................56paucus, Isurus ............108, 110, 114-115, 117-118Pavone ......................................88Pavone di mare ...............................88Péi aspasu ratou ..............................88Peican.....................................107Peix espasa ..................................88Peixe alecrim .................................88Peixe carago .................................96Peixe cavallo .................................88Peixe frade ...................................96Peixe raposo .................................88Peixe rato ....................................88Peixe zorra ...................................88Peje-vaca ....................................96Pelagic thresher ............................81,83pelagicus, Alopias .......................79,81,83pelagicus, Pseudocarcharias .....................72pelagios, Megachasma .........................75pelegrinus, Squalus ............................91Pelerin ......................................96Pélerin ......................................91pennanti, Lamna .............................122pennanti, Squalus .........................121-122pennantii, Selachus ............................91Peregrino .................................91,96Peregrinos ...................................88Peregrinus ...................................90peregrinus, Squalus ............................91perlo, Heptranchias. ............................4peruanus, Gyropleurodus. .......................45peruanus, Heterodontus. ........................46Peruvian horn shark, ...........................46Pesca can. ..................................107Pesce bandiera ...............................88Pesce bannera ................................88Pesce can grande ............................107Pesce cane. .................................107Pesce pavone. ................................88Pesce sorcio. .................................88Pesce volpe ..................................88Pescecane ..................................107Peschanye akuly ..............................56Pescio ratto ..................................88Pesciu can ..................................107Petit taupe ..................................115Pez toro .....................................62Pez zorro ....................................88Philippi ......................................34philippi var. japonicus, Cestracion .............39,48philippi, Cestracion .........................39,42philippi, Lamna ..........................118,122philippi, Squalus ........................34,39,42philippinus, Squalus. ...........................42phymatodes, Chiloscyllium .....................172Pici bistinu ..................................107Pigfish. ......................................45Piked dogfish ..................................5Pintado .....................................208Pintarroja .....................................3Pisce surci ...................................88Pisci cani .........................62,66,107, 125Pisci cuda longa ...............................88Pisci cuduto ..................................88Pisci mastinu ................................107Pisci sciabula turca. ............................88Pisci tunnu ..................................114Piscicani ....................................114Pixi tunnu ....................................96plagiosa, Chiloscyllium indicum var...............174Plagiostoma ...........................31,51,126Plagiostomi. ...........................31,51,126Plagiostomi asterospondyli. ..................51,126Plagiostomi Asterospondyli ......................31Plagiostomi diplospondyli .................31,51,126plagiosum var. interruptum, Scyllium .............174plagiosum, Chiloscyllium ..........158, 165, 173, 175plagiosum, Chiloscyllium indicum var.............174plagiosum, Hemiscyllium. ......................174plagiosum, Scyllium .......................165, 173platensis, Carcharias ...........................58platensis, Odontaspis ........................58,62Plectrosoma .................................108Plectrostoma ................................108Pleurotrema ..............................51,126Pleurotremata. .......................1,31,51,126plunketi, Proscymnodon. .........................4Poisson à volies ...............................96Poisson épée .................................88Pollee maku .................................188Polyprosopus ..............................89-90Polyprosopus macer. ...........................91Polyprosopus rashleighanus .....................90Polyprosopus Rashleighanus .....................90pontoppidani, Halsydrus .....................90-91Poos hee ...................................173Porbeagle .......................3,21,52,121-125Porbeagle shark ..............................125Porbeagles .....................96-97, 118, 123-124Poroderma africanum .........................186Poroderma pantherinum .......................186porosus, Scymnus. ............................196Porpoise shark ...............................114Port Jackson bull-head sharks ....................33Port Jackson shark. ......................20,32,43Port Jackson Shark ............................42Port Jackson sharks .........................33,45portus jacksoni, Squalus. .................32,34,42portusjacksoni, Heterodontus. ................42,45portusjacksoni, Squalus .........................39

Sharks of the World, Vol. 2 263Potrava .....................................188Prionace ....................................103Prionace glauca. ..................99,116,121, 206PRISTIDAE ...................................4PRISTIOPHORIDAE ...........................26PRISTIOPHORIFORMES .................1-3, 16, 26Pristiophoroid .................................27Pristiophoroids ...............................128Proarthri ..................................31-32profundus, Alopias ..........................83,85Proktopterides .........................31,51,126Prosarthri. .................................31-32PROSCYLLIIDAE. ..............................3Proscymnodon plunketi .........................4Pseudocarcharias. ....................57,71-72, 75Pseudocarcharias kamoharai .................71-72Pseudocarcharias pelagicus .....................72PSEUDOCARCHARIIDAE ..............57,71-73. 75PSEUDOCARCHARINIDAE .....................71Pseudoginglymostoma ........129, 185, 188-190, 192,199-200, 202Pseudoginglymostoma brevicaudatum ........199-200Pseudoginglymostomatinae .....................190PSEUDOTRIAKIDAE .........................3,24Pseudotriakis .................................53Psina atlantska ...............................125Psina cavlozuba ..............................114Psina dugonoska .............................114Psina golema .................................96Psina ljudozdera. .............................107Psina zmij ozuba ruzicua ........................66Psina zmijozuba sivka ..........................62Pteromylaeus ................................103punctata, Lamna .........................109, 122punctatum, Chiloscyllium . . 145, 166, 168, 172, 175-176punctatum, Hemiscyllium. ......................175punctatum, Scyllium .......................175-176punctatus, Micristodus. ........................203punctatus, Squalus ............................122punctulatus, Squalus ..........................192Qquinquecarinatum, Scyllium ....................186quinquecornuatum, Scyllia .....................186quoyi, Cestracion ..............................45quoyi, Heterodontus. .....................33,45-46RRa sorrah ...................................173Rabosa ......................................88Radukazame ................................121Raefhajen ....................................88Ragged-tooth .................................66Raggedtooth shark .............................62Ragged-tooth shark .....................3,6,62,66Ragged-tooth sharks ...........................56Raggie ......................................62RAJIDAE ...................................103RAJIFORMES .............................1-2, 16Rajoidei ..................................26,28ramalheira, Gyropleurodus ......................46ramalheira, Heterodontus. ......................46rashleighanus, Polyprosopus. ....................90Rashleighanus, Polyprosopus ....................90rashleighanus, Squalus .........................91Ratfish ....................................1,17Ray.........................................26Ray shark ....................................16Rays ......................................1,27Reef whitetip shark. ...........................198remotus, Carcharhinus ..........................3Renard .................................3,86,88Renard à gros yeux ............................84Renard pélagique ..............................81Renards .....................................78Renards de mer ...............................79Requiem sharks ..................16,27,56,61,103Requim .....................................125Requin .......................................3Requin aveugle des roches .....................145Requin aveugle gris-bleu .......................147Requin baleine ...............................203Requin blanc ................................107Requin crocodile. ..............................72Requin de Port Jackson .........................33Requin dormeur à crête .........................38Requin dormeur bouledogue .....................45Requin dormeur buffle ..........................41Requin dormeur chabot .........................47Requin dormeur cornu ..........................36Requin dormeur d’Oman ........................50Requin dormeur nekozame ......................39Requin dormeur taureau ........................43Requin dormeur zebre ..........................48Requin féroce .................................64Requin grande gueule ..........................76Requin long nez ..............................125Requin lutin ..................................69Requin noronhai ...............................66Requin nourrice ..............................193Requin nourrice fauve .........................197Requin sable ...............................6,62Requin sable tachete ...........................62Requin taureau. ...............................59Requin taureau bamba. .........................62Requin zèbre ................................186Requin-bleu ..................................97Requin-carpette à collarette .....................137Requin-carpette á moustache ...................133Requin-carpette beige .........................141Requin-carpette chat ..........................135Requin-carpette chin ..........................134Requin-carpette roux ..........................138

264 FAO Species Catalogue for Fishery Purposes No. 1Requin-carpette tacheté ........................139Requin-chabot á taches blanches ................174Requin-chabot à taches bleues ..................175Requin-chabot bambou ........................175Requin-chabot birman .........................169Requin-chabot camot ..........................167Requin-chabot élégant .........................172Requin-chabot épaulette .......................180Requin-chabot gris ............................170Requin-chabot grivelé .........................179Requin-chabot indonésien ......................171Requin-chabot marquéterie .....................183Requin-chabot moine ..........................182Requin-chabot ocellé ..........................181Requin-nourrice à queue courte. .................200Requins aveugles. ............................142Requins baleine ..............................201Requins crocodile. .............................71Requins de sable ..............................55Requins dormeurs .............................33Requins grande gueule .........................74Requins lutin. .................................68Requins nourrices ............................188Requins pelerin ...............................88Requins taupe ................................96Requins zébres ..............................184Requins-carpettes ............................130Requins-chabot ..............................164Requins-tapis ............................126, 148Requin-tapis barbu ............................151Requin-tapis cordonnier ........................163Requin-tapis moustache. .......................154Requin-tapis paste ............................158Requin-tapis savetier ..........................160Requin-tapis sombre ..........................161Requin-tapis tacheté ..........................156Requin-taupe commun .........................122Requin-taupe saumon .........................119Reusenhaai ..................................96Reuzenhaal ..................................96Rhinchodon. .............................202-203Rhincodon. ..........103, 126-129, 185, 190, 201-203Rhincodon typus ................23,77,94,202-203RHINCODONTIDAE ........19,127-129, 190, 201-202Rhincodontinae ..............................201Rhincodontoidea .............................126Rhincodontoidei ..............................126Rhinecodon ..............................202-203Rhineodon. ..............................202-203Rhineodon typicus ............................203Rhineodon typus. .............................203Rhineodontiana ..............................201RHINEODONTIDAE .......................201-202Rhineodontini ................................201Rhineodontis .............................201-202Rhiniodon ...............................201-203Rhiniodon typus ..........................202-203RHINIODONTIDAE ....................128, 201-202RHINOBATIDAE. .............................103RHINOBATIDAE ...............................3Rhinobatos ..................................103rhinoceros, Squalus ............................91Rhinocodon. .............................202-203RHINOCODONTIDAE .....................128, 196Rhinodon. ...............................202-203Rhinodon typicus .............................203Rhinodontes .................................201RHINODONTIDAE ....................128, 201-202Rhinodontinae ...............................201Rhinodontini .................................201RHINODONTOIDAE ......................128, 201Rhinodontoidei ...............................201Rhipidion ....................................26Rhizoprionodon ..............................103RHYNCHOBATIDAE ..........................103Rhynchobatus. ...............................103Ridge back shark .............................173Ridgeback catshark ...........................173Ridgebacked bamboo shark, ....................173Riesenhai ....................................96Rineodon. ...............................202-203Rinquim ....................................114River sharks ...................................4Rogatye akuly. ................................33Romano .....................................88Româo ......................................88rondeletii, Carcharoden .........................99rondeletii, Carcharodon .....................99-100rondeletti, Carcharias .........................100rostratus, Cetorhinus ........................89-90rostratus, Somniosus (Rhinoscymnus) ...............2rostratus, Squalus .............................91Rough sharks .................................27Roussette .....................................3Roussette ferrugineuse ........................199rueppelli, Ginglymostoma ......................196russellianum, Chiloscyllium. ....................176russellianus, Squalus (Scyliorhinus) ..........166, 176Rusty carpet shark ........................138-139Rusty catshark ...........................139, 199Rusty shark .................................199Ry brigde ....................................96Rymer. ......................................96SSaddled carpet shark ..........................135Saddled catshark .............................136Sail fish. .....................................96Sailfish ......................................96Sakezame ..................................121Salmon shark ...............20,52,97,119,121, 125Salmon sharks ........................97,118,120

Sharks of the World, Vol. 2 265Salproig ....................................107Salproix ....................................107Salroig ..................................66,107Sand shark ...........................3,6,62,195Sand sharks ................................3,56Sand tiger ................................62,198Sand tiger shark . . 3, 6, 15, 52, 56, 59-60, 62, 66, 103, 198,206Sand tiger sharks .................21,53,55-56, 103Sand tigers ...................................52Sarda ...................................62,107Sardo ......................................125Saw shark. ...................................26Sawfish. ................................4,16,27Sawfishes .....................................1Sawsharks. ...........................2,15-16, 27Sazaewari. ...................................40Sazaiwari ....................................40SCAPANORHYNCHIDAE .......................68Scapanorhynchoidea ...........................51Scapanorhynchus ...........................68-69Scapanorhynchus dofleini .......................69Scapanorhynchus jordani .......................69Scapanorhynchus lewisi. ........................68Scapanorhynchus mitsukurii .....................69Scapanorhynchus owstoni .......................69Scapasaurus ..................................90School shark. ..................................6Scoliophis. ...................................90Scoliophis atlanticus ...........................91Scylia griseum ...............................171Scyliorhinid ..................................22Scyliorhinid catsharks. .........................196SCYLIORHINIDAE ..................3,103, 128, 130Scyliorhinus. ..................................3Scyliorhinus barbatu ..........................155Scyllia ......................................126Scyllia quinquecornuatum ......................186SCYLLIFORMES .............................126Scylliodei ................................51,126Scyllioidei ...............................126, 128SCYLLIOLAMNIDAE ...................51,126, 128Scylliorhininae ...............................128SCYLLIORHINOIDAE .................130, 164, 184Scyllium ....................................152Scyllium barbatus ............................155Scyllium cirratum. ............................192Scyllium cirrhosum ...........................192Scyllium ferrugineum. .........................196Scyllium freycineti ............................179Scyllium griseum. .........................170-171Scyllium heptagonum. .........................186Scyllium malaianum. ..........................196Scyllium malaisianum .........................179Scyllium ocellatum. ...........................181Scyllium ornatum .........................158, 174Scyllium pantherinum .........................186Scyllium plagiosum .......................165, 173Scyllium plagiosum var. interruptum ..............174Scyllium punctatum. .......................175-176Scyllium quinquecarinatum .....................186Scymnodalatias ...............................73Scymnus porosus .............................196Seaape .....................................88Seafox......................................88Sea tiger ....................................188Seaur talay ..................................188seftoni, Heterodontus ...........................42Sel devye akuly ...............................97Selache ...................................88-90Selache elephas ...............................91Selache maxima ...............................91Selache maximum .............................91Selache maximus ..............................91SELACHIDAE. ................................88Selachii .......................1,31-32, 51, 53, 126Selachina .................................88-89Selachio gigante. ..............................96Selachus ..................................89-90Selachus pennantii .............................91Selanche .....................................90Selanonius ..................................117Selanonius walkeri ........................117,122selanonus, Squalus. ...........................122Sevengill shark .................................4Sevengill sharks ...............................20Sevengills ....................................60Shark with tiger-like spots ......................188Sharkray .....................................16Sharks .......................................1Sharp nosed .................................114Sharpnose mako .............................114Sharpnosed .................................114Sharp-nosed mackerel shark ....................114Shavianus ....................................90Shavianus, Squalus (Cetorhinus) ..................91Shima-nekozame ..............................49Shimazame .................................170Shinvala ....................................188Shiroboshi-tenjiku. ............................175Shirowani ....................................62Shortfin mako. ....52,77,97,102-103, 110-111, 113-114Shortfin mako sharks ...........................52Shortfin makos .........................52-53, 103Shortfinned mako shark ........................114Shortnose chimaeras ..........................103Shorttail nurse shark ..........................201Short-tail nurse shark ..........................200Short-tailed nurse shark ........................200Shovelnose shark. .............................62Shovel-nosed shark ............................62Sicklefin lemon shark ..........................198

266 FAO Species Catalogue for Fishery Purposes No. 1Sildehaaen ..................................125Sillhaj ......................................125Silver shark. ..................................17Silver sharks ..............................1-3, 27Singe de mer .................................88Sixgill shark .................................101Sixgill sharks .................................20Skate .......................................16Skates ...................................26,29Slasher ......................................88Sleeper sharks ................................27Sleepy shark. ............................199, 201Slender bamboo shark .....................172-173Smalltooth sand tiger ...........................66Smalltooth sandtiger. ........................53,64Smeriglio ...................................125Smidiru ......................................66smithi, Carcharodon ..........................100smithii, Carcharodon ..........................100Smooth dogfishes. ..............................3Smoothhound ..................................3Smoothhounds .................................3Snapper shark ...............................114Sobraig .....................................114Sobratg .....................................114Solraig ......................................66Solrayo ......................................64Solrayo ojigrande ..............................66Solrayos .....................................55Somniosus (Rhinoscymnus) rostratus ..............2Sorglio pisci tunnu ............................125Soupfin shark ..................................6Southern basking shark .........................96Southern catshark ............................140Southern collared cat shark .....................140spallanzanii, Isurus ...........................109Speckled carpet shark. .....................183-184Speckled cat sharks ...........................177Speckled catshark ............................184Sphyrna ....................................103Sphyrna tiburo ...............................102SPHYRNIDAE ............................56,103Spikkel-skeurtandhaai ..........................62Spiny dogfish .............................27,103Spiny dogfishes ................................3Spiny sharks. .................................27Spitting shark ................................199Spookfish ....................................17Spotted catshark .............................176Spotted ragged-tooth shark ......................62Spotted ragged-tooth sharks ....................113Spotted wobbegong ...................156-157, 163Springhaai ..................................107Squali ................................31,51,126SQUALIDAE .....3,17,32-33, 55, 78, 96, 103, 148, 201SQUALIFORMES ..........................1-3, 27Squalini. ..............................31,51,126Squaliolus. ...................................73Squalo bianco. ...............................107Squalo elefante ...............................96Squalo feroce .................................66Squalo massimo. ..............................96Squaloid .....................................27Squaloid sharks ...............................22Squaloidea ...................................31Squaloidei. .............................31,51,53Squaloids .......................26,29,32,53,128Squalomorph sharks ................1,23,27-28, 128Squalomorphi .................................2Squalomorphii ...........................1,16,27Squalus ..................32,34,88,90,99,103, 117Squalus (Carcharhinus) lamia ...................100Squalus (Carcharias) maou .....................100Squalus (Carcharias) vulgaris. ..................100Squalus (Carcharias) vulpes .....................80Squalus (Cetorhinus) Gunneri ....................91Squalus (Cetorhinus) Shavianus ..................91Squalus (Lamna) cepedii .......................109Squalus (Scyliorhinus) dentatus. .................172Squalus (Scyliorhinus) russellianus. ..............176Squalus (Scyliorhinus) tuberculatus ..............172Squalus (Scyliorhinus) unicolor. .................166Squalus (Scyliorhinus) variegatus ...............166Squalus (Scyliorhinus) waddii ...................145Squalus acanthias ..............................5Squalus alopecias .............................86Squalus americanus .........................58,62Squalus appendiculatus ........................155Squalus argus. ...............................192Squalus barbatus .........................152, 155Squalus carcharias .........................99-100Squalus caudatus .............................172Squalus cetaceus ..............................91Squalus cirratus ..........................191-192Squalus cirrhatus .............................192Squalus cirrosus. .............................186Squalus colax ............................172-173Squalus cornubicus ....................117,121-122Squalus cornubiensis ..........................122Squalus elephas ...............................91Squalus fasciatus .........................185-186Squalus ferox ..............................63-64Squalus glaucus ...........................99,121Squalus gronovianus ..........................172Squalus gunnerianus ...........................91Squalus homianus .............................91Squalus indicus ......................166, 172-173Squalus isodus ................................91Squalus jacksoni ..............................42Squalus jacksonii ..............................42Squalus littoralis ..............................58Squalus lixa ..................................58

Sharks of the World, Vol. 2 267Squalus lobatus ..........................153, 155Squalus longicaudatus .........................186Squalus macrodous ............................58Squalus maculatus ........................152, 155Squalus maximus ...........................90-91Squalus modestum ............................146Squalus monensis .........................121-122Squalus nasus ...........................117,121Squalus ocellatus ........................177, 181Squalus oculatus .............................181Squalus pantherinus. ..........................186Squalus pelegrinus. ............................91Squalus pennanti. .........................121-122Squalus peregrinus. ............................91Squalus philippi .........................34,39,42Squalus philippinus ............................42Squalus portus jacksoni ..................32,34,42Squalus portusjacksoni .........................39Squalus punctatus ............................122Squalus punctulatus ..........................192Squalus rashleighanus ..........................91Squalus rhinoceros. ............................91Squalus rostratus ..............................91Squalus selanonus ............................122Squalus tigrinus ..........................185-186Squalus tuberculatus ..........................172Squalus tygrinus. .............................186Squalus varius. ...........................185-186Squalus vulpecula .............................86Squalus vulpes .............................86,99Squalus vulpinus ...........................80,86Squalus waddi. .......................143, 145-146SQUARINIFORMES. ...........................16Squatina californica. ...........................37Squatinida ............................51,53,126SQUATINIDAE. .......................27,127, 148SQUATINIFORMES ..................1-2, 18, 21, 27Squatinoid ...................................27Squatinoids. ..............................53,128Stegastoma ..................................185STEGASTOMATIDAE .........................184Stegostoma ..................126-129, 184-186, 202Stegostoma carinatum .........................186Stegostoma fasciatum .....................185-186Stegostoma longicaudatus ......................185Stegostoma tigrinum .......................185-186Stegostoma tigrinum naucum. ...................186Stegostoma tigrinus ...........................186Stegostoma tygrinum ..........................186Stegostoma varium ........................185-186STEGOSTOMATIDAE ...........4,127-129, 184, 196Stegostomatinae .........................128, 184Stegostonea .................................185Stingray .....................................16Stingrays ..............................4,60,103Stone shark .............................173, 175strahani, Hemiscyllium. ...................177, 182Striped bamboo shark .........................175Striped bullhead shark ..........................49Striped cat shark ..............................49Striped shark ................................175Sun fish .....................................96Sunfish ......................................96Suño ........................................46superciliosus, Alopias .................79,82-83, 85Surraig ......................................66Sutorectus. ........................3,149, 162-163Sutorectus tentaculatus ................149, 153, 162Swordfin ....................................125Synchismus. .................................166Synodontaspis .............................56,58Synodontaspis taurus ........................58,62TT’ushak’o ...................................97Tasha......................................175Tabarao ....................................107Tabbigaw ....................................45Taburo .....................................107Taiwan saddled carpet shark ....................134Tanifa ......................................107Taniwha ....................................107Tapicero barbudo .............................151Tapicero japoné ..............................154Tapicero manchado ...........................156Tapicero occidental. ...........................161Tapicero ornamentado .........................158Tapicero zapatilla .............................160Tapicero zapatudo ............................163Tasmanian carpet shark ........................139Tasmanian spotted catshark. ....................139Tasselled wobbegong ......................151-152Taulo. ......................................125Taupe .......................................97Taupe bleu ..................................110Taupe longue aile .............................117Tauró blanc. .................................107taurus, Carcharias. 3, 6, 15, 52, 56-58, 62, 65-66, 70, 73,97, 99, 103, 113, 198, 206taurus, Eugomphodus .....................3,58,62taurus, Odontaspis ........................3,58,62taurus, Synodontaspis .......................58,62taurus, Triglochis ..............................58Tawny nurse shark ........................196-198Tawny shark .................................199Teguzame. ...................................96Tengu (goblin) shark. ...........................71Tenguzame. ...............................71,96Tenjikuzame .................................173tentaculatus, Crossorhinus. .....................162tentaculatus, Orectolobus ......................162tentaculatus, Sutorectus ...............149, 153, 162Tetnoras ..................................89-90

268 FAO Species Catalogue for Fishery Purposes No. 1Tetraoras ....................................90Tetraoras angiona .............................91Tetroras. ..................................89-90Tetroras angiova ............................89-91Tetroras maccoyi ..............................91Thintail thresher ...............................88Thornray .....................................16Thrasher shark ................................88Thrashers ....................................79Thresher shark ..........................83,86,88Thresher sharks ......................21,29,78-79Threshers ...........................52-53, 79, 98Tiburo ......................................107tiburo, Sphyrna ..............................102Tiburón ................................3,62,114Tiburón acebrado .............................186Tiburón ballena. ..........................203, 208Tiburón blanco ...............................107Tiburón bocudo. ...............................76Tiburón ciego de roca. .........................145Tiburón ciego gris .............................147Tiburón cocodrilo ..............................72Tiburón dama ................................208Tiburón de leznas ..............................62Tiburón duende ...............................69Tiburón tamborín ..............................46Tiburón vitaminico ..............................6Tiburones acebrados ..........................184Tiburones ballena .............................201Tiburones bocudos .............................74Tiburones ciegos .............................142Tiburones cocodrilo ............................71Tiburones duende. .............................68Tiburones tapiceros .......................126, 148Tiger ragged-tooth .............................66Tiger shark ...........................62,175, 188tigrinum naucum, Stegostoma ...................186tigrinum, Stegostoma ......................185-186tigrinus, Squalus ..........................185-186tigrinus, Stegostoma. ..........................186tigris africanus, Isurus. ........................109tigris, Carcharias. ............................109Time teller shark ..............................173Tintorera ......................................3Tofusa.....................................208Tollo .........................................3Tommy .....................................107Tope.........................................3Torafuzame. .................................188Toro bacota. ..................................59Toro bambaco. ................................62Toros. .......................................55Touilele boeuf taupe ...........................125Touille ......................................125Touille à l’épée ................................88Triaenodon obesus ............................198TRIAKIDAE ...............................3,103tricuspidatus, Carcharias ....................58,62tricuspidatus, Odontaspis .......................58tricuspitatus, Odontaspis ........................58Triglochide feroce ..............................66Triglochide tauro. ..............................62Triglochidini ..................................55Triglochis ..............................55-56, 58Triglochis taurus ..............................58trispeculare, Chiloscyllium .....................183trispeculare, Hemiscyllium .............143, 177, 183Tropidodus ...................................34Tropidopus ...................................34True sharks. ..................................56Tubarâo baleia ...............................208Tubarâo branco ..............................107Tubarâo frade. ................................96Tubarâo raposo ............................85,88Tubarào zebra ...............................188Tubaroes crocodilos ............................71Tubaroes de areia .............................56Tubaroes de leite .............................189Tubaroes dorminhocos. .........................33tuberculatum, Chiloscyllium ....................166tuberculatus, Chiloscyllium .....................172tuberculatus, Squalus. .........................172tuberculatus, Squalus (Scyliorhinus) ..............172Tuk........................................208TukiTuki ...................................208Tuku.......................................208Tuna shark ..................................107Tunnu palamitu di funnu ........................107Turbarâo dorminhoco de Moçambique. .............48tygrinum, Stegostoma. .........................186tygrinus, Squalus .............................186typicus, Rhineodon. ...........................203typicus, Rhinodon. ............................203typus, Rhincodon ...................77,94,202-203typus, Rhineodon .............................203typus, Rhiniodon ..........................202-203typus, Rhinocodon. ...........................203UUbazame ....................................96Umiwani .....................................62unicolor, Squalus (Scyliorhinus) .................166Uptail ......................................107VVaalhai .......................................6Varied carpet shar ............................140Varied catshark. ..............................140Variegated shark .............................188variegatus, Squalus (Scyliorhinus) ...............166variolatum, Hemiscyllium ..................136, 140variolatum, Parascyllium ..................131, 140varium, Stegostoma .......................185-186

Sharks of the World, Vol. 2 269varius, Squalus ...........................185-186Verdoun .....................................62verus, Carcharias. ............................100Viper sharks ...............................27-29Vitamin shark ..................................6Volpe di mare .................................88vorax Carcharias .............................100vulgaris, Squalus (Carcharias) ..................100Vulpecula .................................78,80Vulpecula marina. ..........................80,86vulpecula, Galeus. .............................86vulpecula, Squalus .............................86VULPECULIDAE ..............................78vulpes, Alopecias ..............................86vulpes, Alopias ................................86vulpes, Carcharias ..........................80,86vulpes, Squalus. ............................86,99vulpes, Squalus (Carcharias). ....................80vulpinus, Alopias ..................79,81-83, 86, 99vulpinus, Squalus ...........................80,86Wwaddi, Brachaelurus ..................143-145, 148waddi, Squalus .......................143, 145-146waddii, Squalus (Scyliorhinus) ..................145walkeri, Selanonius .......................117,122wardi, Orectolobus .......................156, 159Water alligator ................................74Water crocodile. ...............................74Weasel sharks ................................16Wedgefish ...................................16Weisshai. ...................................107Western wobbegong ..........................161Whale .......................................77Whale shark ......23,77,94-95, 103, 127, 203, 207-208Whale sharks ....16,24,29,127-128, 201-202, 204-208Whiptail shark. .............................83,88White death .................................107White death shark ............................107White pointer ................................107White shark 21, 52, 61, 97-99, 101, 103, 107, 113, 120, 125White sharks ............6,52-53, 94, 96, 99, 102-103Whitespotted bamboo shark .................174-175Whitespotted bullhead shark .....................47Whitespotted cat shark. ........................175Whitetip shark. ...............................116whitleyi, Lamna ..........................118,122Winged shark .................................16Winged sharks .................................1Witdoodshaai ................................107Withaai .....................................107Wobbegong ...............................3,157Wobbegong sharks ............................24Wobbegongs. 18, 29, 60, 127-128, 148-149, 152, 156-157Wuia .....................................33-34YYahiya. .....................................199yangi, Carcharias .............................72Yasurizame. .................................208Yellow belly. ..................................62Yellow shark ..................................62Yu belangkas ................................173Yu bodoh ...................................173Yu checkak ..................................188Yu tadek ....................................177Yu tokay ....................................188Yu tokek ....................................188Yu tokele. ...................................173ZZebra bullhead shark ........................33,48Zebra horn shark ..............................49Zebra Port Jackson shark .......................49Zebra shark ............................4,185-188Zebra sharks ......................16,24,128, 184zebra, Centracion. .......................34,39,48zebra, Cestracion ...........................39,48zebra, Heterodontus ........................33,48Zev ........................................132Zorra de mar. .................................88Zorro ..................................85-86, 88Zorro blanco ..................................88Zorro cauda longa .............................88Zorro ojón .................................84-85Zorro olho grande. .............................85Zorro pelagico ................................83Zorro pelágico ................................81Zorros ....................................78-79Zozame .....................................96

FAO Species Catalogue for Fishery Purposes No. 1, Vol. 2ISSN 1020-8682SHARKS OF THE WORLDAN ANNOTATED AND ILLUSTRATED CATALOGUE OFSHARK SPECIES KNOWN TO DATEVolume 2. Bullhead, mackerel and carpet sharks(Heterodontiformes, Lamniformes and Orectolobiformes)

FAO Species Catalogue for Fishery Purposes No. 1, Vol. 2FIR/Cat.1/2SHARKS OF THE WORLDAN ANNOTATED AND ILLUSTRATED CATALOGUE OF SHARKSPECIES KNOWN TO DATEVolume 2Bullhead, mackerel and carpet sharks(Heterodontiformes, Lamniformes and Orectolobiformes)byLeonard J.V. CompagnoShark Research CenterIziko-Museums of Cape TownSouth African MuseumCape TownSouth AfricaFOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONSRome, 2002

ii FAO Species Catalogue for Fishery Purposes No. 1The designations employed and the presentation of material inthis publication do not imply the expression of any opinionwhatsoever on the part of the Food and Agriculture Organizationof the United Nations concerning the legal status of anycountry, territory, city or area or of its authorities, or concerningthe delimitation of its frontiers or boundaries.ISBN 92-5-104543-7All rights reserved. No part of this publication may be reproduced, stored in aretrieval system, or transmitted in any form or by any means, electronic,mechanical, photocopying or otherwise, without the prior permission of thecopyright owner. Applications for such permission, with a statement of thepurpose and extent of the reproduction, should be addressed to the Director,Information Division, Food and Agriculture Organization of the UnitedNations, Viale delle Terme di Caracalla, 00100 Rome, Italy.© FAO 2001

Sharks of the World, Vol. 2iiiPREPARATION OF THIS DOCUMENTThis document was prepared in FAO under a special programme made possible thanks to a generous Trust Fund(GCP/INT/643/JPN) from the Government of Japan. The present publication is the second installment of the updatedversion of the original FAO Catalogue of Sharks of the World (Compagno 1984), and constitutes volume two of the firstnumber in a new series: FAO Species Catalogue for Fishery Purposes. Up until now, FAO Species Catalogues had existedas a far too lengthy series of volumes within FAO Fisheries Synopsis No. 125. But given the importance and size of thatseries and the continuing need for this type of publication in the foreseeable future, the species catalogues have grown outas an independent series starting with the present three-volume work. In order to preserve the continuity of the speciescatalogues, the new series will maintain the trademark orange-coloured cover of the old series as well as most of its format.As the new Catalogue has grown apace with new information and revisions, it is being published as three free-standingvolumes, each with separate pagination, introduction, terminology, systematic sections, glossary, list of species by FAOStatistical Areas, and a dedicated bibliography. This will allow readers to independently use each volume without having toconsult the other volumes for technical terms and measurements or bibliographic purposes, as was the case in the oldcatalogue. We hope that this added flexibility will be received as an improvement.Programme manager: Pere Oliver (FAO, Rome).Scientific and technical editor: Ramón Bonfil (Fisheries Centre, UBC, Vancouver).Technical assistance: Jascha Minow (FAO, Rome).Editorial assistance: Michèle Kautenberger and Nicoletta De Angelis (FAO, Rome).Scientific illustrators: Emanuela D’Antoni, Paolo Lastrico, Oliviero Lidonnici, Pier-Luigi Isola and Miloud Sadeir (FAO,Rome), Leonard J.V. Compagno (Shark Research Center, South African Museum, Iziko - Museums of Cape Town), Mary H.Wagner (US Bureau of Commercial Fisheries, Stanford) and Juan Varela (Madrid).Cover illustration: Emanuela D’Antoni (FAO, Rome).Page composition and indexing: Michèle Kautenberger (FAO, Rome).Digitization of distribution maps: Fabio Carocci (FAO, Rome), and Elena V. Orlova.Compagno, L.J.V.Sharks of the world. An annotated and illustrated catalogue of shark species known to date.Volume 2. Bullhead, mackerel and carpet sharks (Heterodontiformes, Lamniformes andOrectolobiformes).FAO Species Catalogue for Fishery Purposes. No. 1, Vol. 2. Rome, FAO. 2001. 269p.ABSTRACTThis is the second volume of an extensively rewritten, revised, and updated version of the original FAOCatalogue of Sharks of the World. The present volume reviews all 15 families 25 genera and 57 species ofliving bullhead, mackerel and carpet sharks (orders Heterodontiformes, Lamniformes andOrectolobiformes), that is, the non-carcharhinoid galeomorph sharks, including certain well-establishedbut currently undescribed species mainly from Australia. It gives accounts for all orders, families and generaand all keys to taxa are fully illustrated. Information under each species account includes: valid modernnames and original citation of the species (or subspecies); synonyms; the English, French, and SpanishFAO Names for the species; a lateral view and often other useful illustrations; field marks; diagnostic features;distribution, including a GIS map; habitat; biology; size; interest to fisheries and human impact; localnames when available; a remarks section when necessary; and literature. The volume is fully indexed andalso includes sections on terminology and measurements including an extensive glossary, a list of speciesby FAO Statistical Areas, an appendix on shark preservation, and a dedicated bibliography.DistributionAuthorsFAO Fisheries OfficersRegional Fisheries Councils and CommissionsSelector SC

iv FAO Species Catalogue for Fishery Purposes No. 1AcknowledgementsIt is hard to summarize the large number of people that have helped me over the course of preparing the 1984 sharkcatalogue and its current revision; several are no longer living, but I salute the memories of the dead as with the living, andmake no distinction here. I apologize beforehand if I have forgotten anybody, which will inevitably happen.I would especially like to thank W.I. (Bill) Follett, of the California Academy of Sciences, San Francisco, California, forencouragement on my initial checklists and interest in shark systematics many, many years ago. Especial thanks go to fiveof my research colleagues, Shelton P. (Shelly) Applegate (Los Angeles County Museum and Instituto de Geologia, CiudadUniversitaria, Mexico City), J.A.F. (Jack) Garrick (Department of Zoology, Victoria University, Wellington, New Zealand),Stewart (Stew) Springer (National Marine Fisheries Service, USA, and Mote Marine Laboratory, Sarasota, Florida), WarrenC. Freihofer (Division of Systematic Biology, Stanford University, California Academy of Sciences and Tiburon Center forEnvironmental Studies) and Sid F. Cook (Argus-Mariner Consulting Scientists, Portland, Oregon), all of whom wereextremely helpful and inspirational over the years. Special thanks for my wife, Martina Roeleveld (South African Museum)for her support.The 1984 catalogue owed an enormous amount to the untiring and titanic efforts of Dr Walter Fischer, formerly MarineResources Department, FAO Fisheries Division, who inspired me to make it a far more useful work than would have beenmy inclination as a professional systematist. This was not entirely a painless process, but I learned an enormous amountwhile writing it. Special thanks go to Dr Bernard Zahuranec, of the US Office of Naval Research, for providing support for thewriting of the catalogue over three years in the form of a research contract to the writer at the Tiburon Center forEnvironmental Studies, San Francisco State University, Tiburon, California, and for arranging funding for a circumglobalresearch trip in 1982. Thanks also to Dr Samuel E. Gruber (University of Miami) and the American Elasmobranch Society forsupport for the publication of the 1984 catalogue in the United States. Dr Cornelia E. Nauen, formerly of the MarineResources Department, FAO Fisheries Division, was extremely helpful in the organization of the final version of the 1984catalogue, which went through several changes in format during the past five years. Thanks also to the staff of Dr W.Fischer’s Species Identification and Data Programme (SIDP) at FAO for their efforts, particularly for the fine artistic work ofEmanuela D’Antoni, Pier Luigi Isola, Paolo Lastrico, and Oliviero Lidonnici in translating often difficult material from theliterature as well as the writer’s research drawings and maps into the illustrations in this catalogue; and to GiuliaSciarappa-Demuro for typing the final manuscript. Mrs Paula Smith, Ms Barbara Nabors, and Ms Dale White of the TiburonCenter for Environmental Studies helped in the typing and copying of versions of the manuscript and the organization of thebibliography.The present revised Catalogue was made much easier on the writer’s side by powerful personal computers, slide andflatbed scanners, OCR and digital graphics programmes, and electronic communications, which eliminated the need tospend part of the time at Rome to produce the Catalogue. A very large thanks to Dr Ramón Bonfil (Fisheries Centre,University of British Columbia), who served as grand coordinator of production and editor for the Catalogue at SIDP, as wellas peer reviewer, user-friendliness tester, and contributor to the work; a massive task, but from this side not thankless!Thanks to Dr Pere Oliver, Dr Kent Carpenter, Mr Jascha Minow, Ms Michèle Kautenberger, Ms Emanuela D’Antoni,Ms Nicoletta De Angelis and Ms Guilia Sciarappa at SIDP, and Mr Fabio Carocci at FAO, for their outstanding work duringvarious stages of the project that helped to make it possible.Special thanks also to the following: Bobb Schaeffer, Donn Rosen, Gareth Nelson, John Maisey, Gavin Naylor, GuidoDingerkus and Marcelo de Carvalho (American Museum of Natural History, New York); John R. Paxton and Doug Hoese(Australian Museum, Sydney); John E. Randall, Arnold Susumoto, J. Culp and Richard Pyle (Bernice P. Bishop Museum,Honolulu); Robert Lea (California Department of Fish and Game); F. R. Harden Jones, John Stevens, Peter Last, A.J. Rees,T. Carter, Maria Bresîc’ and Justine O’Regan (Division of Fisheries, CSIRO Marine Laboratories, Hobart, Tasmania);Alwynne Wheeler, Peter Whitehead, and Oliver Crimmen (British Museum [Natural History], London, now the NaturalHistory Museum); David Ward (University of London); William D. Eschmeyer, Frank H. Talbot, John McCosker, Tyson R.Roberts, Lillian J. Dempster, Robert P. Dempster, Tomio Iwamoto, Pearl Sonoda, and David Catania (California Academy ofSciences, San Francisco); E.G. Silas, K.K. Appukuttan, and M.E. Rajapandian (Central Marine Fisheries Research Institute,Cochin, India); L.W. Filewood (Department of Agriculture, Stock and Fisheries, Konedobu, Papua-New Guinea); GreggBrett (East London Museum, South Africa); Rob Heijman (Expert Center for Taxonomic Identifications, Amsterdam); IanFergusson (European Shark Research Bureau and Shark Trust, UK); Hideki Nakano (National Research Institute of FarSeas Fisheries, Shimizu); Kazunori Yano (Seikai National Fisheries Research Institute, Ishigaki Tropical Station, FisheriesAgency of Japan); Rainer Zangerl (Department of Geology, Field Museum of Natural History, Chicago); Kazuhiro Nakaya(Hokkaido University); Manfred Fischer (Infotek, Cape Town); E. MacPherson (Instituto del Mar, Barcelona); Dario J. Guitart(Instituto de Oceanologia, Academia de Ciencias de Cuba, Havana); Gordon Hubbell (JAWS International, Miami); M.Stehmann (Institut für Seefischerei, Hamburg); J.L.B. Smith, Margaret Smith, Michael Bruton, Tom Hecht, Phillip C.Heemstra, Elaine Heemstra, Paul Skelton, M. Eric Anderson, Dave Voorfelt, Paul Cowley, Billy Ranchod, MargaretCrampton, Robin Stobbs, Edward Matama, S. Matama, H. Tomlinson, Alexandra Macras and K. Lilley (J.L.B. Smith Instituteof Ichthyology, Shark Research Center of JLBSII, and Rhodes University, Grahamstown, South Africa); Supap Monkolprasitand Prachit Wongrat (Faculty of Fisheries, Kasetsart University, Bangkok); Martha J. Mitchill (Kent Cambridge ScientificInc., Palo Alto, California, and Stanford University); Robert J. Lavenberg, Camm Swift and Jeffery A. Seigel (Los AngelesCounty Museum of Natural History); Myvanwy M. Dick (Museum of Comparative Zoology, Harvard University, Cambridge,Massachusetts); Louis Garibaldi and Don Reed (Marine World, Redwood City, California); G. Victor Morejohn, GregorCailliet, and Dan Varoujean (Moss Landing Marine Laboratories); Dave Ebert (Moss Landing Marine Laboratories, SharkResearch Center in Grahamstown and Cape Town, and US Abalone); Ken Goldman (Virginia Institute of Marine Science,

Sharks of the World, Vol. 2vGloucester Pt, Virginia); Lee B. Hulbert (National Marine Fisheries Service, Juneau, Alaska); Marie-Louise Bauchot andBernard Seret (Museum National d’Histoire Naturelle, Paris); Merry Camhi (Ocean Wildlife Campaign, National AudubonSociety, USA, and IUCN Shark Specialist Group); Beulah Davis, Graeme Charter, Geremy Cliff, Sheldon Dudley, B. Wilsonand P. Mthembu (Natal Sharks Board, Umhlanga Rocks); John Moreland and Andrew Stewart (National Museum of NewZealand, Wellington); Sarah Fowler (Nature Conservation Bureau, UK, and IUCN Shark Specialist Group); Ernst Mikschi(Naturhistorisches Museum, Vienna); Keiichi Matsuura (National Science Museum, Tokyo); Che-Tsung Chen (NationalTaiwan Ocean University); Peter Ng and Kelvin Lim (Zoological Reference Collection, Department of Zoology, NationalUniversity of Singapore); Nadaraj (Nat) Kistnasamy, John Bass, Alan Bowmaker, Elinor Bullen, Rudy van der Elst, JohnWallace and Lynnath Beckley (Oceanographic Research Institute, Durban, South Africa); S. Uchida and Katherine M. Muzik(Okinawa Expo Aquarium); Fritz J. Pfeil (Pfeil Verlag, Munich); Malcolm J. Smale, Graham Ross, Colin Buxton, J. Clarke,M. du Plessis, and W. Mahola (Port Elizabeth Museum and Oceanarium: Bayworld); Mabel Manjaji and Alvin Wong (SabahFisheries, Sabah, Malaysia); Anna Wong (Sabah Museum, Sabah); C.G. Alexander, Robert I. Bowman, MargaretG. Bradbury, Erwin Seibel, and Michael Jocelyn (San Francisco State University); Carl L. Hubbs and Richard H. Rosenblatt(Scripps Institution of Oceanography, La Jolla, California); A.I.L. (Andy) Payne, L. Botha, A. Robertson, A. Badenhorst,C. Augustyn, Marek Lipinski, Geremy David, Herman Oosthuizen, Marc Griffiths, Andrew Penny, R. Cooper, Robin Leslie,Roy Melville-Smith, Rob Tilney, H. Crous, Peter Sims, Captain Derek Krige, and the officers and crew of RV AFRICANA, RVALGOA,RVBENGUELA, and RV SARDINOPS (Sea Fisheries Research Institute/Marine and Coastal Management, CapeTown); Barrie Rose, Graham Brill and the captains, officers and crew of the Irvin and Johnson deepwater bottom trawlingfleet (Sea Fisheries Research Institute and Irvin and Johnson, Cape Town); Steve Campana (Bedford Institute ofOceanography, Dartmouth, Nova Scotia); Leslie W. Knapp (Smithsonian Oceanographic Sorting Center, Washington, DC);Martina A.C. Roeleveld, P.A. (Butch) Hulley, Graham Avery, Sydney Kannemeyer, Rachel Alexander, Mark Marks, AndreaBertolini, Michelle van der Merwe, Liz Hoenson, Cedric Goliath, Mike Bougaardt, Ricardo Adams, Mike Boon, and PeteWhite (Iziko: Museums of Cape Town, South African Museum and Shark Research Center); George S. Myers, NormanK. Wessells, Paul R. Ehrlich, Donald Kennedy, Robert T. Schimke, Evelyn Shaw, John H. Thomas, and Margaret A. Sharp(Department of Biological Sciences, Stanford University); Carlos J. Villavicencio-Garayzar (Universidad Autónoma de BajaCalifornia Sur, La Paz, Mexico); John D. McEachran (Texas A and M University); Takashi Okutani and Hajime Ishihara(Tokyo University of Fisheries); Stanley H. Weitzman and Victor G. Springer (Division of Fishes, US National Museum ofNatural History); John G. Casey, and Harold L. Pratt, Jr. (US National Marine Fisheries Service, Narragansett Laboratory);Bruce B. Collette, Susumu Kato, Joseph Russo, Susan Smith and Mary H. Wagner (US National Marine Fisheries Service);Hans-J. Paepke (Museum für Naturkunde, Universität Humboldt, Berlin); C. Richard Robins and Arthur Myrberg (Institute ofMarine Sciences, University of Miami); Reeve M. Bailey (University of Michigan Museum of Zoology, Ann Arbor); BruceWelton (University of California, Berkeley, Los Angeles County Museum of Natural History, and Chevron Oil Field ResearchCompany); Dave Freer, Richard Brooke, Tim Crowe, Charles Griffiths and J. Jarvis (University of Cape Town); FernándoMárquez (Instituto Nacional de la Pesca, Mexico); Steve Kajiura (University of Hawaii, Honolulu); Tom Thorson (Universityof Nebraska); Toru Taniuchi (University of Tokyo, Department of Fisheries); Alvaro E. Tresierra Aguilar (UniversidadNacional de Trujillo, Peru); Wolf-Ernst Reif (Institut und Museum für Geologie und Paläontologie der Universität Tübingen,Germany); Fabio H. V. Hazin (Universidade Federal Rural de Pernambuco, Recife, Brazil); George Zorzi (Sacramento StateUniversity and US Air Force); R.J. McKay, Gerald Allen, J. Barry Hutchins and Nick Haigh (Western Australian Museum,Perth, Australia); Leighton R. Taylor, Jr. (Waikiki Aquarium, Honolulu, Hawaii); John C. Cordell (Woods Hole OceanographicInstitute, Massachusetts); John W. Shipman (Zoological Data Processing, Socorro, New Mexico); P.K. Talwar (ZoologicalSurvey of India, Calcutta).The writer’s work on both versions of the Catalogue was aided by funding and other support from the Department ofFisheries of FAO and the Fisheries Agency of Japan; the US Office of Naval Research, National Science Foundation (NSF),and National Marine Fisheries Service; the UK Darwin Initiative; Argus-Mariner Consulting Scientists; the South AfricanNational Research Foundation (formerly Foundation for Research Development); the J.L.B. Smith Institute of Ichthyology,Iziko: Museums of Cape Town, South African Museum); Tiburon Center for Environmental Studies, San Francisco StateUniversity; CIAC (Cephalopod International Advisory Council); and the Australian Division of Fisheries, CommonwealthScientific and Industrial Research Organization (CSIRO).

vi FAO Species Catalogue for Fishery Purposes No. 1Table of Contents1. INTRODUCTION..............................................................................11.1 Plan of the Catalogue . .....................................................................21.2 Technical Terms and Measurements...........................................................71.2.1 Picture Guide to External Terminology of Sharks ............................................71.2.2 Picture Guide to Skeletal Terminology of Sharks. ............................................91.2.3 Measurements Used for Sharks........................................................111.2.4 Glossary of Technical Terms . .........................................................152. SYSTEMATIC CATALOGUE . ..................................................................312.1 Order HETERODONTIFORMES - Bullhead sharks...............................................312.1.1 Family HETERODONTIDAE . .........................................................33PageHeterodontus . .....................................................................34Heterodontus francisci . .............................................................36Heterodontus galeatus...............................................................38Heterodontus japonicus..............................................................39Heterodontus mexicanus . ............................................................41Heterodontus portusjacksoni..........................................................42Heterodontus quoyi . ................................................................45Heterodontus ramalheira . ...........................................................46Heterodontus zebra . ................................................................48Heterodontus sp. A . ................................................................492.2 Order LAMNIFORMES - Mackerel sharks . .....................................................512.2.1 Family ODONTASPIDIDAE . ..........................................................55Carcharias........................................................................57Carcharias taurus . .................................................................58Odontaspis........................................................................63Odontaspis ferox . ..................................................................64Odontaspis noronhai . ...............................................................662.2.2 Family MITSUKURINIDAE. ...........................................................68Mitsukurina . ......................................................................69Mitsukurina owstoni . ...............................................................692.2.3 Family PSEUDOCARCHARIIDAE......................................................71Pseudocarcharias . .................................................................72Pseudocarcharias kamoharai . ........................................................722.2.4 Family MEGACHASMIDAE . ..........................................................74Megachasma . .....................................................................75Megachasma pelagios . ..............................................................752.2.5 Family ALOPIIDAE .................................................................78Alopias...........................................................................80Alopias pelagicus. ..................................................................81Alopias superciliosus................................................................83Alopias vulpinus . ..................................................................862.2.6 Family CETORHINIDAE .............................................................88Cetorhinus . .......................................................................90Cetorhinus maximus . ...............................................................91

Sharks of the World, Vol. 2Page2.2.7 Family LAMNIDAE..................................................................96Carcharodon . .....................................................................98Carcharodon carcharias . ...........................................................100Isurus...........................................................................108Isurus oxyrinchus. .................................................................109Isurus paucus.....................................................................115Lamna . .........................................................................117Lamna ditropis....................................................................119Lamna nasus . ....................................................................1212.3 Order ORECTOLOBIFORMES - Carpet sharks.................................................1262.3.1 Family PARASCYLLIIDAE . ..........................................................130Cirrhoscyllium....................................................................132Cirrhoscyllium expolitum . ..........................................................133Cirrhoscyllium formosanum . ........................................................134Cirrhoscyllium japonicum...........................................................135Parascyllium . ....................................................................136Parascyllium collare . ..............................................................137Parascyllium ferrugineum...........................................................138Parascyllium variolatum . ...........................................................140Parascyllium sp. A . ...............................................................1412.3.2 Family BRACHAELURIDAE .........................................................142Brachaelurus . ....................................................................144Brachaelurus waddi. ...............................................................145Heteroscyllium....................................................................146Heteroscyllium colcloughi...........................................................1472.3.3 Family ORECTOLOBIDAE. ..........................................................148Eucrossorhinus . ..................................................................150Eucrossorhinus dasypogon . .........................................................151Orectolobus . .....................................................................152Orectolobus japonicus..............................................................154Orectolobus maculatus . ............................................................155Orectolobus ornatus . ..............................................................158Orectolobus wardi . ................................................................159Orectolobus sp. A . ................................................................161Sutorectus . ......................................................................162Sutorectus tentaculatus . ............................................................1622.3.4 Family HEMISCYLLIIDAE . ..........................................................164Chiloscyllium.....................................................................165Chiloscyllium arabicum. ............................................................167Chiloscyllium burmensis . ...........................................................168Chiloscyllium griseum..............................................................169Chiloscyllium hasselti . .............................................................171Chiloscyllium indicum..............................................................172Chiloscyllium plagiosum . ...........................................................173Chiloscyllium punctatum............................................................175Hemiscyllium.....................................................................177Hemiscyllium freycineti.............................................................179Hemiscyllium hallstromi . ...........................................................180Hemiscyllium ocellatum . ...........................................................181Hemiscyllium strahani..............................................................182Hemiscyllium trispeculare...........................................................183vii

viii FAO Species Catalogue for Fishery Purposes No. 1Page2.3.5 Family STEGOSTOMATIDAE . .......................................................184Stegostoma. ......................................................................185Stegostoma fasciatum . .............................................................1862.3.6 Family GINGLYMOSTOMATIDAE . ....................................................188Ginglymostoma . ..................................................................191Ginglymostoma cirratum............................................................192Nebrius . ........................................................................195Nebrius ferrugineus................................................................196Pseudoginglymostoma..............................................................199Pseudoginglymostoma brevicaudatum . ................................................2002.3.7 Family RHINCODONTIDAE. .........................................................201Rhincodon . ......................................................................202Rhincodon typus . .................................................................2033. LIST OF SPECIES BY MAJOR FISHING AREAS. ..................................................2104. APPENDIX . ...............................................................................2114.1 Keeping Sharks for Scientific Study . .........................................................2114.1.1 Generalities About Shark Collections...................................................2114.1.2 Practical Advice for Creating Shark Collections . ..........................................2125. BIBLIOGRAPHY............................................................................2156. INDEX OF SCIENTIFIC AND VERNACULAR NAMES...............................................249click for next page

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