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Shark Attack versus Ecotourism: Negative and Positive Interactions

A.PeterKlimley Dept.ofWildlife,Fish,andConservationBiology,Universityof,Davis,California TobeyH.Curtis SustainableFisheriesDivision,NationalMarineFisheriesService,Gloucester,Massachusetts ABSTRACT : Unprovokedattacksbyonareexceptionallyrarephenomena.Sharkstypicallyhavetwomotivations, feedingordefense,thatresultinattacksonhumans.Threespecies,thebull,tiger,andwhitesharks,areresponsibleforthemajority ofattacksonhumans.Thesepredominantlyfeeding-motivatedattacksareoftentheresultofthemistakingitsvictim fornaturalprey.Manyspecies,however,exhibitadefensive,aggressivedisplaythat,unheeded,mayresultinasinglebiteor slashingwoundtoahuman.Thenumberofunprovokedattacksbysharksonhumansworldwidehasrisenfrom8during1900- 1904,ofwhich2werefatal,to330during2000-2004,ofwhich29werefatal.Theratesof5.8fatalitiesperyearduring2000-2004 and6.4fatalitiesperyearduring1995-1999arenegligiblerelativetotheaverageof42,593fatalitiesperyearduetoautomobile accidentsreportedfrom1993-1995intheUnitedStatesalone.Takingalookatsharksfromanotherperspective,ecotourismhas becomeimmenselypopularinthe1990sand2000s.Thereareopportunitiestoviewsharksinthewildoneverycontinentexcept Antarctica, with the scalloped hammerhead, white, , and reef sharks being among the most popular subjects. Shark ecotourismisprovidingthepublicwithanobservationalexperiencethatcanbeaspleasurableaswhalewatching,anditcanbea cost-effectivealternativesourceofemploymentforfishermen.Thiscouldleadtoreducedsharkfishingincertainregionsofthe worldandenablesharkpopulationstorecovertotheirformerlevelsofabundance. KEY WORDS : agonisticdisplay,ecotourism,publicsafety,sharkattack Proc.22 nd Vertebr.PestConf. (R.M.TimmandJ.M.O’Brien,Eds.) PublishedatUniv.ofCalif.,Davis. 2006. Pp.33-44. INTRODUCTION beach recreation is characteristically at its peak, were Sharksaremanyoftheoceans’toppredators.Along reportedbythemediawithgreatsensation,andthisbad withrays,sharkscomprisethesubclassElasmobranchii,a pressresultedinbathersfromtheNewJerseyandNew groupofmostlycarnivorous,cartilaginousfishesthatare York metropolitan areas avoiding their customary presentinalloftheworld’soceans.Eventhoughtheover recreationaldestinationsfortherestofthesummer.This 403speciesofsharksmakeuponly1.4%ofthe27,977 notoriousseriousofevents,whichadverselyaffectedthe species of fishes inthe oceans (Nelson 2006), none are economy of many coastal communities in New Jersey, morefearedbyhumanswhoentertheocean.Atleast30 wasthereal-lifeeventwhichwasthebasisofthefictional species of sharks have been confirmed attackers of accountinPeterBenchley’s . humans,butonlyafewofthelargestpredatoryspecies, ThisphenomenonisnotconfinedtotheUnitedStates the white ( Carcharodon carcharias ), tiger (Galeocerdo ofAmerica.A16-yearoldboy,MichaelHely,wasbitin cuvieri ), and bull sharks ( Carcharhinus leucas ), are thehipbyaragged-toothedshark( Carchariastaurus )at responsible for the majority of attacks (Caldicott et al . Amazintoti, Natal during April 1960 (Davies and 2001). D’Aubrey 1961 a), and although not fatal, this shark Asharkattackonahumanusuallyattractsimmediate attack was reported widely in the South African press. and extensive attention by the media. The newspaper Thiswasfollowedbyafatalattackona25-yearoldmale, headlines and articles reporting the attack are usually PetrusSithold,138kmawayatMargate,amajorresort sensational and frightening and may have a negative towninNatal,duringDecemberofthesameyear.These impact on tourism in coastal recreational areas. The attacks,andtwoadditionalonesonyoungboys,13and magnitudeoftheimpactisamplifiedgreatlyifmorethan 15yearsold,on6and22ofJanuary1961(Daviesand oneattackoccursatroughlythesameplaceatthesame D’Aubrey1961 b,c)created widespreadfearinthelocal time. For example, Charles Vansant was killed by a community. The news of these attacks was reported shark swimming in the waters near Beach Haven, New widely in newspapers across and resulted in a Jersey,on1July1916.Fivedayslater,on6July,Charles greatreductioninthenumberoftouristsvisitingDurban, BruderwaskilledbyasharkatSpringLake,NewJersey, afavoritedestination,overthenext5years.Theneedto 70 km from Beach Haven. Just 6 days following that learn more about sharks’ motivations to attack humans, attackon12July,asimilardistanceaway,twomenwere and to develop ways of avoiding it, led in part to the fatallyattacked,andonemanwasseriouslymauledbya creation of the South African Association for Marine sharkatMatawanCreek,NewJersey(SchultzandMalin Biological Research and the building of the Durban 1975, Fernicola 2001). Though the species responsible Aquarium,andeventuallyledtothesettingofprotective for these attacks has not been conclusively determined, nets along the of South to exclude sharks most experts suggest that the attacker(s) was either a fromthebeachesandthusreducetheriskofsharkattack white shark or a bull shark. These 5 attacks, which (CliffandDudley1991). occurred close to the 4 th of July holiday, when public Overthelast46yearssincetheattacksinDurban,the 33 publicperceptionofsharkshasslowlychangedfromone withover80%ofreportedattacksfollowingthatpattern offearandmisunderstandingtooneofmorefascination. (Caldicott etal .2001).Swimmersandsurfersinshallow Thischangeinpublicopinionhasbeencausedinpartby waters are common victims of this type of attack, in theavailabilityofmoreaccuratescientificknowledgeof whichtheshark makesasinglestrike,leavingabiteor the behavior and ecology of sharks (e.g., Gilbert 1966, slash wound before retreating and not returning. The Hodgson and Mathewson 1976, Klimley and Ainley victimrarelyseestheattackerduringthebriefinteraction, 1996, Carrier et al . 2005). This research effort was andthewoundsaregenerallyminor.Ananalysisof86 stimulatedatfirstbytheNavy’sinterestinidentifyingthe South African shark attacks showed that 81% of attack motivation(s) for sharks to attack humans, in order to victims suffered only minor injuries, requiring only avoid attacks or deter them while in progress. Even primarysutures(Woolgar etal .2001). thoughtheoccurrenceofsharkattackshasrisenslowlyas Victimsof“sneak”and“bumpandbite”attackstend human utilization of the seas has risen, sharks are now to suffer more severe injuries. These attacks tend to beingperceivedbymanytobevaluablemarineresources, occurindeeperwatersondiversorswimmersandmay bothasafoodsourceandamagnetforecotourism,and involvemultiplestrikes.Thesharkisnotseenpriortothe notsomuchasdangerouspests. “sneak”attack,whichtendstobeapowerfulandviolent The purpose of this article is to review both the interaction typical of a predatory attempt by the shark negativeandpositiveaspectsoftherelationshipbetween usinganambushtactic.In“bumpandbite”attacks,the sharksandhumans.Wewillfirstidentifythemotivations shark circles the victims, often bumping them with its that lead sharks to attack humans. Our next step is to snoutpriortoattack.Thisbehaviorisbelievedtoallow establishhowcommonlysharksattackhumansandhow the shark to assess the palatability of the potential prey significant the risk of losing one’s life due to a shark item, or to determine the potential threats it may pose attack is, relative to losing one’s life due to other more priortomakingadecisiveattack.Alargepercentageof commonhazards.Wethenbrieflydiscussthedeleterious these types of attacks result in fatality (Caldicott et al . effectofhumanfisheriesonsharkpopulations.Thenext 2001). subjectdiscussedissharkecotourism,apastimeenjoyed by divers worldwide that come to view sharks in their MotivationtoAttackHumans own habitat, and we discuss attempts to convert One can avoid being attacked if one decreases the fishermen,whoheretoforekilledsharks,intomembersof motivationofthesharktoattack.Hungerisoneobvious theecotourismcommunity. motivationforasharkattacktoattackahuman.Attack wounds often do resemble bites by which the shark METHODS attemptedtofeed.Ontheotherhand,themagnitudeand We base our discussion of the motivation of shark shapeofattackwoundsdo notalwaysbearthisout.If attacksonareviewofthescientificliterature.Thestatis- attacksarefeedingmotivated,onewouldexpecttheprey ticsonattacksonhumansbysharksarebasedonrecords tobeconsumedeitherpartiallyorwholly.Indeed,thisis fromtheInternationalSharkAttackFile(ISAF),whichis thecaseinsomeinstances.DuringOctober,1939,two maintainedattheMuseumofNaturalHistoryat diverswerewitnessedbeingattackedoffabeachinthe theUniversityofFlorida.TheISAFisanarchiveofall province of New South Wales, , and on the knownsharkattacksworldwide.Sinceitsestablishment following day their remains were recovered from the in1958,morethan4,000individualshark-humaninterac- stomach of a 3.5-m (total length) tiger shark (Klimley tion investigations have been compiled, covering the 1974).Oftentheevidenceisnotsoconclusive;remains period from the mid-1500s to the present. Unless of victims have been found in shark stomachs, but the otherwise noted, all references to shark attacks only attackshavenotbeenwitnessed.Thevictimsmaywell include cases of “unprovoked” attack, defined as have drowned before being consumed, and hence are incidents where an attack on a live human by a shark classifiedasscavengingeventratherthananattack. occursinitsnaturalhabitat,withouthumanprovocation In many instances, victims of shark attacks have of the shark (G. Burgess, ISAF, pers. commun.). We experiencedthelossofonlysmallamountsofflesh.One obtained the statistics on deaths due to various injuries such example occurred off West Palm Beach in 1968 from the National Safety Council, and these were (Klimley 1974). A young boy was attacked close to averaged over 3 years, 1993-1995. The compilation of shore while snorkeling. Teeth marks in the shape of a opportunitiesforsharkecotourismwereprimarilybased half-moonwereimpresseduponthebottomofbothofhis on The Shark Watchers Handbook: A Guide to Sharks swimming fins. These marks could only result from a and Where to See Them (Carwardine and Watterson slashingmovementmadebyoutstretchedjawsratherthan 2002). averticalbite.Severewoundswereinflictedbetweenthe boy’skneeandankle.Thesealsolackedthecharacteris- RESULTS ticpuncturewoundsofthepointedgrippingteethofthe Thecauseandendresultofasharkattackwillvary shark’slowerjaw.Althoughcloseto1,000sutureswere dependingonthelocation,theactivityandresponseofthe neededtoclosethechild’swounds,verylittlefleshwas victim, the species involved, and the motivation behind lost. This observation led a U.S. naval physician to its attack. Caldicott et al . (2001) describe 3 general publish an important scientific article, entitled “Shark categories of shark attacks: 1) “hit and run” attacks, 2) Attack:FeedingorFighting,”questioningthetraditional “sneak”attacks,and3)“bumpandbite”attacks.“Hitand hunger-motivatedexplanationandsuggestingthatattack run”attacksarebyfarthemostcommontypeofattack, behaviormightratherbedefensiveinnature,triggeredby 34 intrusion into a protected area by the victim (Baldridge andWilliams1969). Defense Agonistic displays are conspicuous and exaggerated motor patterns that demonstrate the displaying individual’silleaseduetothepresenceofanotherandits capacity to inflict harm, should the intruder come any closer(Burghardt1970).Thehunchingposture,erection of hair, baring of teeth, and hissing of a cat when confronted by a barking dog, for example, is a display conveyingtothedogthecat’sreadinesstodefenditself byattacking,ifthedog movesanycloser.Thedisplay enablesboththecatanddogtoavoidtheinjuriesthatthey wouldsustaininafight. Suchadisplaywasphotographedandanalyzedforthe grayreefshark, Carcharhinusamblyrhynchos (Nelson et al. 1986).Itsdisplaywasbrokendowninto2ambulatory elements and 4 postural elements, for the sake of understandingitbetter.Theformerareveryexaggerated tail movements on a horizontal plane and swimming to the side in a looping configuration (Figure 1). The 4 postural components are the upward pointing of the snout,loweringofthepectoralfins,archingoftheback, and bending of the tail in a lateral direction (Figure 2). The intensity of the display, evinced by the degree of compression of the looping trajectory of the shark, increases with the speed at which the diver approaches the shark and the degree of the shark’s confinement (Figure3).Thedisplayresultsinanattack,ifanintruder doesnotretreatbutcontinuestoapproachtheshark.This relationshipwasdemonstratedexperimentallybysteering asubmersibleofthesizeofalargesharktowardagrey reef shark, eliciting an agonistic display, yet failing to Figure2.A.Photographofgrayreefsharkfromasideview retreat but continuing to approach the shark. The performingagonisticdisplay,evidentbyslightlyopened reactions of the displaying sharks were to accelerate mouth,upwardpointingsnout,depressedpectoralfins, explosively upward, around, and downward to inflict a archedback,andlateralbendofposteriorbodyandtail; slashingbiteonthesideofthesubmersible(Nelson etal. B.Viewfromposteriorofsharkswimminginrolling 1986). Similar displays have been observed by the mode;andC.Sideviewofnormallyswimmingshark blacknose (Carcharhinus acronotus ), lemon (Negaprion (takenfromJohnsonandNelson1973). brevirostris ), silky (Carcharhinus falciformis ), and bonnetheadsharks(Sphyrnatiburo )(MyrbergandGruber 1974;Klimley,unpubl.observations).

Figure3.Diagramofagonisticdisplaywithswimmer Figure1.Diagramofagonisticdisplay:A.Topandfrontal approachingsharkswimmingatincreasingswimming viewofnormalswimming;B.Similarviewsofdisplaywith speeds.Notecompressedloopingtrajectoryexhibited laterallyexaggeratedbeatingofcaudalfin;C.Sideand towardrapidlyapproachingdiver.Thishighintensity frontalviewsofspiralloopingtrajectoryofshark (taken displayisoftenfollowedbythesharkstrikingthediver fromJohnsonandNelson1973). (takenfromJohnsonandNelson1973).

35 Predation shortdurationsweredefinedaseventsandrepresentedby Though humans are not a major component of any alphabetical codes. Continuous actions with longer shark’s normal diet, there are 3 species of sharks that durationsweredefinedasstatesandwererepresentedas havebeendocumentedtopreyonhumansonoccasion. horizontal bars. Most of the behavioral patterns were They are the bull shark, the tiger shark, and the white directlyobservedacts,suchasthepreybeingseizedby shark.Adulttigersharksfeedonadiversedietoflarge thesharkwhileeitherinahorizontalorverticalposition such as bony fishes, sharks and rays, birds, (V).Afewbehavioralpatternswereindirectindicatorsof andsea(Lowe etal .1996),andhumanshavebeen the sharks seizing the prey, such as the sudden appear- foundintheirstomachs.Adultbullsharksoccasionally ance of blood, or splashing at the anterior of the shark prey on marine mammals including sea (Klimley, (SA).Ananalysisofthesediagramssuggestedthatpre- unpubl. observation) and (Heithaus 2001), and datorybehaviorofthewhitesharkcanbebrokendown they are fully capable of consuming humans. This intoasequenceoftypicalbehavioralstatesandactions. species enters estuaries and rivers and was possibly In eastern Pacific waters, white sharks feed most responsiblefortheattackson3bathersinMatawanCreek frequentlyonnorthernelephantseals( Miroungaangusti- inJuly1916.Itisdifficulttoconcludethatwhitesharks rostris ), like those that live on the shores of the South that attack humans off Central California, southwestern FarallonIslands.Thesequenceofbehaviorsinpredation coastofAustralia,andsoutherncoastofAfricaintendto P-120 is an example of a white shark feeding bout in feed on humans (as argued by Burgess and Callahan whichthepreyisaseal.Observerswerealertedtothe 1996),becausethehumanmaybeconfusedwithnormal shark’sinitialstrikebytheappearanceofasmallbright- prey, seals and sea lions, which are more abundant at redareaontheseasurface(eventB).Thesharksurfaced theselocations(TricasandMcCosker1984). and began to swim with exaggerated tail beats 0.3 min Because divers and surfers on the Pacific coast of later,possiblywiththesealheldinitsjaws(stateSWX). NorthAmericahopetoavoidattacksbywhitesharksand Thesealsurfacedafter1.5minandfloatedmotionless,no hopetosurviveattackswhentheyhappen,agreatdealof longer bleeding, 75-100 m from the site of the initial attentionhasbeendevotedinthepopularmediatowhat strike (state MOT). The seal was no longer inside the happenswhenasharkattacks.Dothewhitesharkseat areatintedredwithblood;theareahadbynowexpanded people,orjustbitethem?Doesashark“biteandspit”– and elongated toward the site where the dead seal thatis,expelitspreyandreturnlatertoeatit,expectingit reappeared.Thesealhadalreadylostamassiveamount tobleedtodeath?Cananattackingsharkbebeatenoff ofblood.Thecarcassremainedatthesurfacefor1.1min with a stick? Why do sharks carry their prey for long beforethesharkseizedthecarcassandsubmergedwithit. periods of time before taking a bite? Observations of Splashingandasmallamountofbloodwereobserved1 natural white shark predation at the South Farallon minlaterneartheheadoftheshark,indicatingthatthe Islands, California, have provided possible answers to sealhadbeenbittenagain(eventsSAandB).Later,the manyofthesequestions(Klimley etal .1996).Detailed carcass was seized 4 more times by the shark from diagramswereconstructedof129videotapedpredations underneath,withnolossofblood(eventV).Duringthe between 1989 and 1992. These video records were feedingbout,thesealmovedadistanceof133.4malmost transcribedintodiagrams.Shownisadiagramofawhite directlyoffshoreofthelocationwheretheattackbegan. shark predation on a seal (Figure 4). Each behavioral Theintervalbetweentheappearanceofblood(eventB) patternofthesharkandpinnipedwasrecordedaseither andthebloodlessandimmobilesealatthesurface(state an“event”ora“state”(Lehner1979).Discreteactionsof MOT) was 1.5 min, almost twice the duration of the

Figure4.Thesequenceofbehavioraleventsandstatesinafeedingboutofawhitesharkonaseal.PredatoryattackP- 120onanorthernelephantseal,15October1989.Theactivityofthewhitesharkisgivenabovetheabscissa;thatofthe prey,below.Behavioraleventsareindicatedbyalphabeticalcodesabovethelinesleadingfromtheabscissa. Key:B=blood-stainedwater,SA=splashatanteriortorsoofshark,V=verticalsharkbite.Behavioralstatesaredenotedbyhorizontalbars. SWS(solidbar)=sharkswimmingslowly,SWX(cross-hatchedbar)=sharkswimmingwithexaggeratedtailbeats,MOT(clearbar)=sealfloating motionlessatsurfaceofwater.

36 0.8-min interval between the second and third events (eventV,stateCAR,andeventR).Whilebeingcarried (onsetofstateMOTandeventSA),andalmost3times inthesharksjaws,thevictimlostmuchblood(eventB) thedurationofthe0.6-minintervalbetweenthethirdand (R.Lea,pers.commun.).Similartothesituationduring fourthevents(eventsSAandV). feeding bouts on pinnipeds, the shark carried the diver Thebehaviorofthesharkandsealcanbeinterpreted underwaterforsomedistance.Thediverstrucktheshark in the following way, based on the review of this and with the butt of a bang stick 3 times before the shark many other predations. The initial strike was rarely releasedhim(eventsSTandR)(R.Lea,pers.commun.). witnessedduringanattack,suggestingthatthepreywas This pattern is consistent with the notion that the shark seizedunderwater.Rather,theobserversweremostoften exsanguinatesitsprey,butthesharkreleaseditshuman alerted of the attack by a spot of a large blood-stained prey intact and swam off. In a seal attack, the shark areaofwater.Thebloodstainelongatedinonedirection, wouldhavechaseddownthepreyandfinisheditoff.The andthenthesharkreappearedwiththesealbesideit.In pattern Tisserand reported, in which he was bitten and other attacks, the shark would be seen swimming with then released, is typical of white shark encounters with wide tail beats. Such beats would be necessary for the people off California (Tricas and McCosker 1984). In sharktopropelitselfforwardifitwerecarryingtheheavy another attack, a shark seized and released a brown seal in its jaws. This sequence of events appears to pelican,eventhoughitwasquicklydisabledandunable indicatethatthesharkcarriesitspreybeneaththesurface to resist further attack. The bird was left bleeding and then returns to the surface. After a prolonged profusely and struggling spasmodically at the surfaced interval,thesealthenfloatstothesurfaceinanimmobile untilitdied2minuteslater.Thesharkneverreturnedto statewithabiteoffleshmissing,although,significantly, feedonthepelican(Klimley etal .1996).Furthermore, thewoundsinmostcaseswerenolongerbleeding.The many sea otters are found whole but dead along the sharkthensurfacedquicklyandswamtothecarcassand California coast, with fragments of white-shark teeth seizedit.Thisscenarioisconsistentwiththesuggestion embeddedintheiropenwounds(Ames etal .1996).A that the white shark kills its prey by exsanguination, or seaotterhas yettobefoundinthestomachofawhite blood deprivation. It appears the shark holds the seal shark(Klimley1985). tightlyinitsjawsuntilitisnolongerbleeding. There may be a connection between these observa- Acommonlyheldbeliefisthatsharksbiteseals,and tions.Humans,birds,andseaottersarecomposedmainly thenquicklyspitthemoutinawoundedbutintactstateto of muscle, whereas the preferred prey of sharks– seals die,beforeagainattackingthem.Thesharkschasedand and cetaceans (dolphins and )– are composed captured64%ofthesealsthatsurvivedtheoriginalattack mainly of fat. Sharks may prefer energy-rich marine attheFarallones.Furthermore,thesharksdidnotreturn mammals to other comparatively energy-deficient spe- immediately to feed on the carcass once the seal died. cies.Supportingthisisthepropensityofwhitesharksto Sealcarcasseswereobservedfloatinginimmobilestates feedselectivelyontheblubberofbaleenwhalesandnot foraslongas140minutesbeforesharksreturnedtofeed the tissue underneath (Curtis et al . 2006). This may onthem. explain why white sharks rarely completely consume Interestingcontrastsoccurintheresponsesofsharks humans when they attack. Finally, experiments have to different prey. At 2 p.m. on September 9, 1989, a shownthatwhitesharksdofeedonthefatflensedfrom sharkattackedahuman,andtheattackbeganlikemany seals,butnotontheremainingmuscleonthebodyofthe attacksonseals(Figure5).Awhitesharkseizedtheleg seal(Klimley1994). of Mark Tisserand, a commercial abalone diver, while Furtherevidenceofthereluctanceofwhitesharksto Tisserandwas5-8metersbelowthesurface,nearly200 consumehumanswhollycomesfromtheISAF.Inonly metersfromshore,pausinginapronepositiontoclearhis 6.2%oftheattacksrecordedworldwidefrom1910-1995 ears (Robert Lea, pers. commun.). According to the didawhitesharkremove3bitesfromahuman(Burgess victim (Mark Tisserand, pers. commun.), “the shark and Callahan 1996), and it is unlikely that the humans swamupfromunderneath,seizedme,carriedmedown werecompletelyeateninthisnumberofbites.Twobites for5to7seconds,andsuddenlyletmegoandswamoff” were observed in 16.8% of the attacks. In striking

Figure5.Sequenceofattacksonahumanwhowasnoteaten.AttackP-99onahumanon9September1989. Key:V=verticalsharkbite,R=sharkreleasesitem,ST=preystrikesshark

37 contrast,only1seaescapedfromapredatorywhite 350 100 shark during the 129 attacks on pinnipeds recorded by TotalAttacks TotalFatalities 90 300 video at the Farallon Islands, resulting in a predatory OverallFatalityRate(%) successrateof99.2%(Klimley,unpubl.data).However, 80 it is possible that some of these attacked seals escaped 250 70 60 fromthesharksandwereunseenafterwards.Asuccess 200 rate of 55% was recorded for a larger sample size of 50 150 predations(2,088overaperiodfrom1997-2003)occur- 40 ring within an hour of sunrise at Seal Island, South 100 30 NumberofAttacks Africa, and predation ceased later in the day, when the 20

50 PercentageofAttacksFatal successratedecreasedto40%(Martin etal .2005).Inthe 10 majorityofattacksonhumans,56.8%ofthe125recorded attacks worldwide, the shark bit only once and then 0 0 9 9 14 19 34 39 94 904 909 9 9 924 929 9 9 944 949 954 95 964 97 984 989 9 999 004 departed; no bites were removed from the human in 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 0- 5- 0- 5- 0- 0- 5- 0- 5- 0- 5- 0- 5- 5- 0- 0 92 94 96 98 98 00 10.4%oftheattacks.Finally,innoneofthesharkattack 190 19 191 191 1 1925-193 193 1 1945-1950-195 1 1965-19691970-1974197 1 1 1990-1199 2 cases were the intact remains of the whole human recoveredfromthestomachoftheattackingsharkafter Figure6.Frequenciesofnon-fatalandfatalattacksand itscapture(Burgess,pers.commun.).Yet,thepossibility percentagesofthetotalnumberofattacksthatwerefatal existsthatahighlyhunger-motivatedwhitesharkmight during5-yearperiodsshownforallspeciesofsharkson consumeahumanintheabsenceofmorenutritiousprey. humansfrom1900to2005.

Thebullandtigersharksmaynotneedtooptimally 45000 forage on energy-rich prey as does the white shark. 40000 Thesespeciesareectothermic–theirbodytemperatureis 35000 equaltothatofthesurroundingwater,andhencetendto 30000 occupywarmtropicalwaters.Onthecontrary,thewhite 25000 shark, which occupies temperate and polar waters, is 20000 endothermic– maintaining its body temperature above 15000 thatofthecoldsurroundingwaters(Goldman1997).It 10000 5000 possesses multiple retia mirabilia, vascular counter- 0 currentheatexchangers,towarmitsvisceratoaccelerate HumanFatalities(peryear) Motor Falls Drowning Drowningin Lightning the digestion of prey and warm its brain, ensuring Vehicles during Bathtub uniformintegrativeprocessing,anditseyes,tospeedup Recreation theprocessofvisualprocessing(BlockandCarey1985). TypesofAccidents Thisisalsoanenergeticallycostlymetabolicstyleoflife, and it may require that endotherms such as the white 250 shark prey on high-energy prey (Block and Finnerty 200 1994). 150 NegativeInteractions AttacksonHumans 100 The number of attacks of all species of sharks 50 worldwideonhumansper5-yearperiodhasrisenfrom8 0 during1900-1904,ofwhich2werefatal,to330during HumanFatalities(peryear) Horses Hornets, Dogs(USA) Sharks White 2000-2004, of which 29 were fatal (Figure 6). The (USA) Wasps, (Worldwide) Sharks number has risen steadily, and this is likely due to the Bees(USA) (Worldwide) continuousincreaseinthepopulationofhumansworld- wideandtheincreasedfrequencyofhumanutilizationof AnimalsCausingDeaths the ocean. The observed increase in attacks is also, in Figure7.Otherriskstohumans: (above)averageratesof part,anartifactoftheincreasedreportingefficiencyofthe fatalitiesduetoaccidentsintheUnitedStatesover3-year ISAFoverthelast20years,includingtheexpandeduse period,1993-1996; (below)averageratesoffatalitiesdue of the Internet and communication with cooperating tointeractionsofotherspeciescomparedwiththerates scientific observers around the world. Yet, the average ofsharkattack. yearlynumberofattacksbyallsharksduring2000-2004 was only 66 attacks per year, of which 5.8 attacks per recent years is the result of vastly improved medical year were fatal. This was slightly greater than the 266 response and treatment for traumatic injuries like those totalattacks,54.2attacksperyear,and6.4fatalitiesper receivedfromsharkbites. yearduetoattacksrecordedfortheprior5years,1995- The risk of being attacked by a shark is negligible 1999.Twelveand9%ofthetotalnumberofattackswere relativetootherrisksencounteredbyhumansonadaily fatal during the 1995-1999 and 2000-2004 periods, basis.Comparetherateof5.8fatalitiesperyearduring respectively. Thus, comparatively few of the attacks 1990-1994totheaverageyearlyratesoffatalitiescaused werefatal,ascomparedtothe60%fatalityratefromthe by other everyday risks during a 3-year period from 1905-1909period.Thelowpercentageoffatalattacksin 1993-1995(Figure7).Thenumberoffatalitiesinmotor 38 vehicle accidents within the U.S. averaged 42,593 per acttoreducethelikelihoodofbeingattacked.Itisim- yearoverthis3-yearperiod.Thedeathsduetoafall,a perative not to continue swimming toward the shark, common cause of the loss of life among the elderly, particularlyatarapidrate,ortoforceitintoconfinement averaged13,524peryear.Manymorepeoplediefrom suchasagainstthefaceofacoralorrockyreef.When these two causes in the United States than from shark confined,thesharknolongerhastheoptionofflightand attacks worldwide. Furthermore, the number of islikelytofight. Itslikelihoodofdoingsoisapparent worldwide fatalities due to shark attack worldwide was from the speed at which it swims, and the compressed considerably lower than the deaths due to drowning nature of its trajectory. The shark likely will not during recreation (averaging 791 deaths per year), accelerate directly toward the diver’s torso to deliver a drowninginabathtub(296deathsperyear),ordyingby bite,butwilldashbytheperson’ssideandquicklyturn beinghitbyaboltoflightning(72deathsperyear)inthe aroundtoseizeanotherpartofthebody,thearmorleg, U.S.Morecommonweredeathsfromfallingoffahorse and inflict a bite, which usually does not result in the (220 deaths per year) or from being stung by a hornet, removaloffleshbutaslashingwound. wasp, or bee (49 deaths per year) in the U.S. Finally, Ifoneseesalargepredatorysharksuchasabull,tiger, therearefewerdeathsfromattacksbypacksofdogsin orwhiteshark,itisimportanttofacethesharktoindicate the U.S. (16 deaths per year) than deaths due to shark that you are aware of its presence. Predators often attacksworldwide. eschewattackinganimalsthatareawareoftheirpresence Amorerobustwaytoaddresstherelativeriskofshark becausetheyriskinjuryfromthedefensivecapabilitiesof attack is to compare it to the risk of other hazards theprey,suchasbeingbittenbytheteethorscratchedby associated with ocean beaches. Accident records from their claws. If accompanied by a buddy, it is best to theUnitedStatesLifesavingAssociation(USLA)forthe move so that you are back to back and can thus year 2000 compiled by the ISAF provide a more clear collectivelyseeinalldirections.Itisessentialthatyoudo measureofthechanceofbeingattackedbyasharkwhen notappearfrightenedandtrytorapidlyswimawayfrom atanoceanbeach(ISAF,unpubl.data,see:www.flmnh the predatory shark, as this “withdrawal”-type behavior .ufl.edu/fish/Sharks/Statistics/beachattacks.htm). From mayelicitastrikefrompredators. 68 U.S. Atlantic and Pacific lifesaving jurisdictions, beachattendance(264.2million)wasrecordedaswellas PositiveInteractions the number of medical responses (236,642), rescues SharkFisheriesandTheirCollapse (70,771), drownings (74), and other beach-related Humanshavebenefitedattimesinthepastbyfishing fatalities(58).TheISAFcomparedthesestatisticstothe forsharks,butthesefisherieshaveoftencollapsedover numberofsharkattacks(23)reportedfromthesame68 time,leavingsharkfishermenbereftofemployment.We beach areas in 2000. None of these 23 shark attacks eat the flesh of sharks, macerate their fins and use the resultedinafatality.Basedonthesenumbers,theodds resulting colloid to thicken soups, grind their of suffering a drowning or other beach-related fatality cartilaginous skeletons ground into a powder and pill were1in2million,whereastheriskofbeingattackedby (chondrichthin), which when consumed is believed to a shark was 1 in 11.5 million, and the odds of a shark strengthen the cartilaginous lining of the human bone inflictedfatalitywere0in264.2million.Therefore,there joints,andfabricatebeltsmadefromtheirnon-elasticskin areotherpotentialhazardsmorelikelytobeencountered, covered with dermal denticles. However, this taxon even when attending a beach area where shark attack possesseslifehistorycharacteristicsthatmakeitsensitive couldbeapossibility. tooverfishing. For example, basking-shark ( Cetorhinus maximus ) AvoidingSharkAttack fisherieshavecollapsedinboththeAtlanticandPacific Toavoidbeingattackedbyashark,itisbesttotake Oceans(Camhi etal .1998).Averylocalizedfisheryfor some basic precautions. Such precautions include thespeciesaroseinwesternIrelandin1947nearAchill avoiding areas where sharks may be feeding and their Island. Between 900 and 1,800 of these enormous preyisabundant(e.g.,sealcolonies,nearschoolsoffish), plankton-eating sharks were captured each year from andactinginamannerthatmaymimictheirnaturalprey. 1950to1956;overthefollowingyearsthecatchdeclined, Donotenterthewaterifthereischum(maceratedfish) shrinking to 119 in 1968. The fate of the soupfin dispensed during fishing activities. Do not enter the (Galeorhinus galeus ), a relative of the reef sharks, is water alone, as many sharks tend to target solitary anotherprimeexample(Camhi etal .1998).Theneedfor individuals. Avoid murky waters or swimming in low high-grade oil by the military during World War II lightconditionswhenashark’svisionislimited.Ifyou created a market for soupfin liver oil. The liver, areinthepresenceofashark,itisobviouslyprudentnot impregnatedwithoil,canreachathirdofthebodymass to harass or otherwise threaten it, and remove yourself ofanadultshark.Thepriceofoilrosefrom$50perton from the water as quickly and calmly as possible in1937to$2,000pertonin1941.Thecatchofsoupfin (Caldicott etal .2001). rosefrom270tonsperyearintheearly1930stoapeak When confronted by a threatening shark, it may be 2,172tonsin1941andthendroppedto287tonsin1994. helpfulrecognizethemotivationoftheattacker–whether Commercialfisheriesforsharkshaveoperatedinthe itisattackinginordertodefenditselfortofeeduponyou. United States since the 1930s, but they were originally For that reason, it is good to wear underwater goggles smallandrestrictedtosmallareas.Theydidnotbeginto wheninwaterswheresharksarecommon.Hence,one grow until the late 1980s (see state-by-state statistics in canrecognizetheagonisticdisplayofashark,andthen Camhi et al . 1998). Massachusetts, until very recently, 39 had the largest fishery for sharks in the United States. managementof39speciesin3categories:largecoastal This fishery mainly targeted the spiny dogfish ( Squalus sharks,smallcoastalsharks,andpelagicsharks.Quotas acanthias ),whichlivesinlargeschoolsoffNewEngland havebeensetforthecommercialfisheriesandbaglimits inthespringandsummerandthenmigratestothewaters for the recreational fisheries for the large coastal and offthesoutheasternUnitedStatesduringwinter(Jensen pelagic sharks, to lessen the fishing pressure for these 1966). Historically considered to be a valueless, pest species.Commercialfishersarerequiredtoholdafederal species,itisnowserved as“fishand chips”in permittofishforsharksandtoreporttheirlandingsby restaurantsinboththeU.S.andEurope.Thelandingsof category.ThislatterregulationpermitsNMFStomonitor thissmallbutveryabundantpredatoryspeciesroseuntil catch per unit effort on an annual basis and regulate the late 1990s before leveling out, despite increased fishingpressurebaseduponknowledgeofwhethercatch fishingeffort.IntheNationalMarineFisheriesService is increasing or decreasing annually, relative to the (NMFS) 2003 stock assessment for spiny dogfish, the estimatedstockabundance.Finally,theplansprohibitthe stockwasclassifiedasoverfished,despiteregulationsthat wastefulpracticeof“finning”,bywhichonlythefinsof were implemented in 2000 that eliminated all directed sharksareretainedandtherestofthebodyisdiscarded. dogfish fisheries in the U.S. Atlantic (NMFS 2003). Excludingdogfishfromthecatch,Floridahashistorically Ecotourism:ViewinginNaturalHabitat had the largest shark fishery off North America. The The perception that all sharks attack humans was fisherytargetsmanycoastalspeciesintheGulfofMexico dispelledintheearly1980sbyscientificstudies(Klimley and Atlantic waters, primarily sandbar ( Carcharhinus 1981, 1982; Klimley and Nelson 1981), which were plumbeus )andblacktip( C.limbatus )sharks.Theland- featuredinnaturedocumentaries,inwhichscientistswere ingsofthesereefsharksroseuntiltheearly1990s,when seen swimming among schools of hammerhead sharks they also began to decrease, despite increased fishing without being attacked by the members of the schools. effort.Thegrowthinthecommercialsharkfisheriesin Hammerhead sharks were considered at that time to be the late 1980s for sharks in the Atlantic and Gulf of thethirdmostdangerousspeciesofshark(Klimley1974). Mexicowaspartlyduetoanewpublicappreciationfor Scalloped hammerhead sharks feed primarily on fishes the value of these species for food. However, a more andsquid(Klimley1987)andthusmightnotbeexpected importantreasonfortheexpansionofthesharkfisheries tofeedonhumans.Thisisincontrasttothewhite(Tricas was the demand in for highly-prized fins (NMFS and McCosker 1984, Klimley 1985), tiger (Lowe et al . 1993). 1996),andbullsharks(Heithaus2001),whichdofeedon Therearemultiplereasonswhysharksareparticularly mammalian prey and might be expected to attack vulnerabletointensefishingpressure(HoenigandGruber humans.Itissafertoswimamongschoolsofpiscivorous 1990,Klimley1999).Mostsharkspeciesarenearthetop sharks,whicharenotasaggressiveandsolelyconcerned of the food chain and are not abundant. Sharks grow with maintaining their uniform separation distance and slowly,maturelateintheirlives,donotreproduceevery commonheadingwithotherschoolmembers(Nelson et year, and have few young. Take, for example, the al . 1986), than confronting a solitary shark, which may sandbar and scalloped hammerhead sharks ( Sphyrna exhibit an agonistic display when frightened by an spp.), two species frequently caught in the Atlantic and intruder when confined by a nearby reef (Johnson and GulfofMexicofisheries.Thesandbartakes12-15years Nelson1973). toreachmaturityandthengivesbirthto6-10pupsevery Shark ecotourism has become immensely popular in otheryear(ColletteandKlein-MacPhee2002).Female the 1990s and 2000s, and has become worldwide in its hammerheadscantakeupto15yearstoreachmaturity scale. There are opportunities to view sharks in North, andthengivebirthto12-40pups,every1-2years(Camhi Central, and South America, the Caribbean Islands, etal .1998).Thespinydogfishhasalonggrowthperiod Europe,Asia,Africa,andAsia.Thediversityofspecies toreproductivematurity(6-12years)andproduces1-15 thatmaybeviewedunderwaterisgreat,andthesespecies youngeverysecondyear,afteran18-22monthgestation areidentifiedtogetherwiththecountryandlocalityofthe period(ColletteandKlein-MacPhee2002).TheAtlantic ecotourismindustryinTable1.Diversfromalloverthe cod,bycontrast,reachesreproductivematurityinonly2- worldnowtakerecreationalexpeditionstoseamountsand 3 years, produces 1-12 million eggs, and reproduces islands inhabited by schools of hammerhead sharks, in every other year (Collette and Klein-MacPhee 2002). ordertoviewsharksintheirownhabitat.Theschoolsof Thus,forsharksthereisalowerratiobetweenthenumber sharksavoidSCUBAdiversbecausetheyarefrightened of young produced and the number of breeding adults, bythehissingsoundsemittedasairisdrawnthroughthe and if a substantial number of adults are captured and regulator, or by the pulsed, low frequency sounds and consumed, the population can not be expected to be reflection of light associated with the bubbles of air replenished. oscillating back and forth as they rise to the surface By the end of 1980s, concern was voiced over the (Klimley and Nelson 1981). Most sharks can be unmanaged expansion of shark fisheries in the Atlantic approachedmorecloselybymakingbreath-holdingdives Ocean, considering the vulnerability of the species for or using “rebreathers” that re-circulate air with the overfishing(HoenigandGruber1990).In1989,NMFS additionofoxygenandavoidtheproductionofbubbles. begantodevelopamanagementplanforthesharksofthe Therearediveoperationswithboatsthattakeecotourists Atlantic Ocean and Gulf of Mexico. The plan was toviewschoolsofhammerheadsharksatseamountsoff implementedin1993astheFisheryManagementPlanfor thesoutheasterncoastoftheBajaPeninsulaintheGulfof Sharks of the Atlantic Ocean, and is directed at the California,theRevilligigedosIslandsoffMexico,Cocos 40 IslandsoffCostaRica,MapeloIslandoffColumbia,and Herpetologists for permission to use the figures from Johnson and Wolf and Darwin Islands of the Galapagos Islands off Nelson(1973). Ecuador. Publicviewingofwhitesharksfromasharkcagehas LITERATURECITED alsobecome verypopular. Sharkscanbe viewedfrom AMES ,J. A., J. J. GEIBEL ,F. E. WENEDELL ,AND C. A. PATTISON . the safety of a shark cage, yet they must usually be 1996. White shark-inflicted wounds of sea otters in attracted with a mixture of fish body fluids, oils, and California,1968-1992.Pp.309-316 in :A.P.Klimleyand maceratedtissues,called“chum”.Thereareexpeditions D.G. Ainley(Eds.),GreatWhite Sharks:TheBiologyof toviewwhitesharksattheSouthFarallonIslandsoffthe Carcharodoncarcharias .AcademicPress,SanDiego,CA. coastofCentralCaliforniaintheU.S.,GuadalupeIsland BALDRIDGE , J. R., AND J. WILLIAMS . 1969. Shark attack: offthePacificcoastofBajaCalifornia,Mexico,multiple feedingorfighting?MilitaryMedicine134:130-133. islands off the southern coast of South Africa, and BLOCK ,B. A., AND F. G. CAREY .1985.Warmbrainandeye DangerousReefandtheNeptuneIslandsinSpencerGulf temperaturesinsharks.J.Comp.Physiol.B156:229-236. offthesoutherncoastofAustralia.Thereisconcernthat BLOCK , B. A., AND J. R. FINNERTY . 1994. Endothermy in food given to these sharks, which are attracted for fishes: a phylogenetic analysis of constraints, predisposi- viewing, is a positive reinforcement, and trains them to tions,andselectionpressures.Envir.Biol.Fishes40:283- approachanyboat,regardlessofwhethersharkdivingis 302. carriedaboardit.Hence,ecotourismoperatorsarepro- BURGESS , G. H., AND M. CALLAHAN . 1996. Worldwide hibited from feeding sharks once they arrive at tourist patternsofwhitesharkattacksonhumans.Pp.457-469 in : boatsinSouthAfrica.Feedingofsharksdoesoccurat A.P.KlimleyandD.G.Ainley(Eds.),GreatWhiteSharks: Guadalupe Island and in Spencer Gulf. The likelihood TheBiologyof Carcharodoncarcharias .AcademicPress, that sharks learning to associate boats with the SanDiego,CA. opportunity to feed is dependent on the frequency with BURGHART ,G.M.1970.Defining“communication”.Pp.5- whichthesamesharksareattractedandfeednearaboat. 18 in : W. H. Johnston, Jr., D. G. Moulton, and A. Turk InSpencerGulf,lessthan5%ofthesharksviewedfrom (Eds.), Communication by Chemical Signals. New protectivecageshavebeenobservedtoreturn5ormore Appleton-Century-Crofts,NewYork,NY. times,thebelievedthresholdformakingtheassociation CALDICOTT ,D. G. E., R. MAHAJANI ,AND M. KUHN .2001.The betweenfoodandaboat(Robbins2005). anatomyofasharkattack:acasereportandreviewofthe Sharkssuchasreef( Carcharhinus spp.)andhammer- literature.Injury(Int.J.oftheCareoftheInjured)32:445- headsharkscanbeviewedyearroundinsubtropicaland 453. tropical latitudes. However, many of these species CAMHI ,M.1998.Sharksontheline:astate-by-stateanalysis migrate into the higher latitudes during the summer of sharks and their fisheries. Living Oceans Program, months and can be viewed at this time in temperate NationalAudubonSociety,Islip,NY. waters.AtsomelocationsintheCaribbeanandtheSouth CAMHI , M., S. FOWLER , J. MUSICK , A. BRÄUTIGAM , AND S. Pacific,reefsharksareattractedandhand-fedbydivers. FORDHAM .1998.Sharksandtheirrelatives:ecologyand Whale sharks can often be viewed at fish spawning conservation. Occasional Paper #20, IUCN Species aggregationsitesatavarietyoftropicallocations,where SurvivalComm.Int.UnionConserv.NatureNat.Resour. they gather to feed (Table 1). Divers can observe the (IUCN),Gland,Switzerland. behaviorsharksatthesesitesandlearnfirst-handthatthe CARRIER ,J., J. MUSICK ,AND M. HEITHAUS .2005.Biologyof bulkofthesharkspeciesareoflittledangertohumans, SharksandTheirRelatives.CRCPress,BocaRaton,FL. and this has helped to dispel their much-maligned 608pp. reputation. It has also helped to promote greater CARWARDINE , M., AND K. WATTERSON . 2002. The Shark conservationofmanythreatenedsharkspeciesaroundthe WatchersHandbook:AGuidetoSharksandWheretoSee world. Tourists should always, however, keep in mind Them.PrincetonUniversityPress,Princeton,NJ.232pp. that sharks are potentially dangerous predators, and CLIFF , G., AND S. F. J. DUDLEY .1991.Sharkscaughtinthe should always be treated with care when in their protective gill nets off Natal, South Africa. 4. The bull environment. Shark ecotourism is providing the public shark Carcharhinusleucas Valenciennes.S.Afr.J.Marine with an observational experience that can be as Sci.10:253-270. pleasurable as whale watching, and can be a cost- COLLETTE , B. B., AND G. KLEIN -MAC PHEE (E DITORS ).2002. effectivealternativesourceofemploymentforfishermen. BigelowandSchroeder’sFishesoftheGulfofMaine,3 rd Thiscouldleadtoreducedsharkfishingincertainregions Ed.HarperCollins,NewYork,NY.882pp. of the world, and thereby enable shark populations to CURTIS ,T. H., J. T. KELLY ,K. L. MENARD ,R. K. LAROCHE ,R. recovertotheirformerlevelsofabundance. E. JONES ,AND A. P. KLIMLEY .2006.Observationsonthe behaviorofwhitesharksscavengingfromawhalecarcass ACKNOWLEDGEMENTS atPointReyes,California.Calif.FishandGame93:113- WewouldliketothankGeorgeBurgessandAlexiaMorganofthe 124. FloridaMuseumofNaturalHistoryforaccesstosharkattackstatistics DAVIS ,D. H., AND J. D. D’A UBREY .1961 a.Sharkattackoff from the International Shark Attack File, which is supported by the theeastcoastofSouthAfricaon24December,1960with AmericanElasmobranchSociety.FurtherinformationontheISAFcan notes on the species of shark responsible for the attack. be found on the web at: http://www.flmnh.ufl.edu/fish/Sharks/ Investigational Report #2, Oceanographic Research sharks.htm.WealsothanktheAmericanSocietyofIchthyologistsand Institute,Durban,SouthAfrica.

41 DAVIS ,D. H., AND J. D. D’A UBREY .1961 b.Sharkattackoff KLIMLEY , A. P., P. PYLE , AND S. D. ANDERSON . 1996. The the east coast of South Africa, 6 January, 1961. behaviorofwhitesharkandpreyduringpredatoryattacks. Investigational Report #3, Oceanographic Research Pp. 175-191 in : A. P. Klimley and D. G. Ainley (Eds.), Institute,Durban,SouthAfrica. Great White Sharks: The Biology of Carcharodon DAVIS ,D. H., AND J. D. D’A UBREY .1961 c.Sharkattackoff carcharias .AcademicPress,SanDiego,CA. nd the east coast of South Africa, 22 January, 1961. LEHNER , P. N. 1979. Handbook of Ethological Methods. Investigational Report #4, Oceanographic Research GarlandSTPMPress,NewYork,NY.403pp. Institute,Durban,SouthAfrica. LOWE , C. G., B. M. WETHERBEE , G. L CROW , AND A. L. FERNICOLA ,R. G.2001.TwelveDaysofTerror:ADefinitive TESTER . 1996.Ontogeneticshiftsandfeedingbehaviorof Investigationofthe1916NewJerseySharkAttacks.The the tiger shark, Galeocerdo cuvier , in Hawaiian waters. LyonsPress,Guilford,CT.400pp. Environ.Biol.Fishes47:203-212. GILBERT ,P.W.1963.SharksandSurvival.D.C.Heath& MARTIN , R. A., N. HAMMERSCHLAG , R. C. COLLIER , AND C. Co.,Boston,MA.578pp. FALLOWS . 2005. Predatory behaviour of white sharks GOLDMAN ,K.J.1997.Regulationofbodytemperatureinthe (Carcharodoncarcharias )atSealIsland,SouthAfrica.J. whiteshark, Carcharodoncarcharias .J.Comp.Physiol.B MarineBiol.Assoc.U.K.85:1121-1135. 167(6):423-429. MYRBERG ,A. A., AND S. H. GRUBER . 1974.Thebehaviorof HEITHAUS , M. R. 2001. Predator-prey and competitive the bonnethead shark, Sphyrna tiburo . Copeia 1974:358- interactions between sharks (order Selachii) and dolphins 374. (orderOdontoceti):areview.J.Zool.(Lond.)253:53-68. NMFS (NATIONAL MARINE FISHERIES SERVICE ). 1993. HODGSON , E. S., AND R. F. MATHEWSON . 1978. Sensory FisherymanagementplanforsharksoftheAtlanticOcean. BiologyOfSharks,Skates,andRays.OfficeofNavalRes., U.S.Dept.ofCommerce,NOAA,NMFS.167pp. th Dept.oftheNavy,Arlington,VA.U.S.Govt.PrintingOff. NMFS (NATIONAL MARINE FISHERIES SERVICE ). 2003. 37 666pp. Northeast regional stock assessment workshop advisory HOENIG ,J. M., AND S. H. GRUBER .1990.Lifehistorypatterns report. Northeast Fisheries Science Center Reference in the elasmobranches: implications for fisheries manage- Document03-17,U.S.Dept.ofCommerce,NOAA,NMFS. ment. Pp. 1-16 in : H. L. Pratt Jr., S. H. Gruber, and T. NELSON , D. R., R. R. JOHNSON , J. N. MCKIBBEN , AND G. G. Taniuchi (Eds.), Elasmobranchs as living resources: PITTENGER . 1986. Agonistic attacks on divers and advancesinthebiology,ecology,systematics,andstatusof submersibles by gray reef sharks, Carcharhinus thefisheries.NOAATech.Rep.NMFS90. amblyrhynchos :antipredatoryofcompetitive?Bull.Marine JENSEN ,A.C.1966.Lifehistoryofthespinydogfish.Fish. Sci.38:68-88. Bull.654:527-554. ROBBINS ,R. 2004.Environmentalfactorsaffectingsexualand JOHNSON ,R. H., AND D. R. NELSON . 1973.Agonisticdisplayin size segregation, and the effect of baiting on the natural the gray reef shark, Carcharhinus menisorrah , and its behaviours,ofgreatwhitesharks, Carchardoncarcharias , relationshiptoattacksonman.Copeia1973:76-84. at the Neptune Islands, South Australia. Ph.D. dissert., KLIMLEY , A. P. 1974. An inquiry into the causes of shark Univ.ofTechnology,Sydney,Australia. attack.SeaFrontiers20:66-75. SCHULTZ , L. P., AND M. H. MALIN . 1975. A list of shark KLIMLEY , A. P. 1981. Grouping behavior in the scalloped attacksfortheworld.Pp.529-551 in :P.W.Gilbert(Ed.), hammerhead.Oceanus24:65-71. SharksandSurvival.D.C.HeathandCo.Lexington,MA. KLIMLEY , A. P. 1982. Social organization of schools of TRICAS ,T. C., AND J. E. MCCOSKER .1984.Predatorybehavior scallopedhammerheadshark, Sphyrnalewini (Griffithand ofthewhiteshark( Carcharodoncarcharias ),withnoteson Smith), in the Gulf of California. Ph.D. dissert., Univ. itsbiology.Proc.Calif.Acad.Sci.43:221-238. Calif.-SanDiego. WOOLGAR , J. D., G. CLIFF , R. NAIR , H. HAFEZ , AND J. V. KLIMLEY ,A. P. 1985.Thearealdistributionandautoecology ROBBS . 2001. Shark attack: review of 86 consecutive of the white shark, Carcharodon carcharias , off the west cases.J.Trauma:Injury,Infect.Crit.Care50(5):887-891. coastofNorthAmerica.So.Calif.Acad.Sci.,Memoirs9: 15-40. KLIMLEY ,A. P. 1987.Thedeterminantsofsexualsegregation inthescallopedhammerhead, Sphyrnalewini .Envir. Biol. Fishes18:27-40. KLIMLEY , A. P. 1994. The predatory behavior of the white shark.Am.Scient.82:122-133. KLIMLEY , A. P. 1999. Sharksbeware. Am. Scient.87:488- 491. KLIMLEY , A. P., AND D. G. AINLEY (E DITORS ).1996.Great White Sharks: The Biology of Carcharodon carcharias . AcademicPress,SanDiego,CA.517pp. KLIMLEY , A. P., AND D. R. NELSON . 1981. Schooling of scalloped hammerhead, Sphyrna lewini , in the Gulf of California.Fish.Bull.79:356-360.

42 Table1.Region,country,province,andlocalityofvariousecotourismopportunitiesworldwidewithlistofthosesharksthatcanbeviewedateachsite.

Province Species Region Country Locality /State AG BL BK HM HR LM LP MK NR RF ST SV SX SM TH SW TG WB WL WH ZB NorthAmerica Canada BritishColumbia HornbyIsland,BarkeleySound UnitedStates RhodeIsland PointJudith NorthCarolina Papoose,Caribsea,andAtlasWrecks 2 Florida Semantha'sLedge,TenecoTowers,PalmBeach 5 FlowerGardenBanks 2 Washington SeattleSeacreastPark FarallonIslands California ElkhornSlough SanMiguel,SantaCatalina,SanClementeIslands Maui,Oahu,MidwayAtoll 4 Mexico YucatanPeninsula QuintanaRoo 2 BajaCalifornia GuadalupeIsland,ElBajoSeamounts,GordaBank 8 Central Belize BelizeBarrierReef TurneffeAtoll,Glover'sReef,GladdenSpit 3 America Honduras CaribbeanCoast BayIslands 2 CostaRica PacificCoast CatalinaIslands,CanoIsland CocosIslands AlcyoneSeamount,ManuelitaIsland 4 Caribbean Bahamas Bahamas Walker'sCay,GrandBahama,Bimini 7 Islands Turksand TurksandCaicos WestCaicos,FrenchCay,SaltCay 2 3 Caicos Dominican Dominican CatalinitaandSaonaIslands 2 Republic Republic Cuba Cuba PlayaSantaLucia,Jucaro 4 Cayman CaymanIslands BloodyBayMarinePark,GrandCayman 3 Islands Trinidadand Trinidadand FlyingReef,Diver'sDream 3 Tobago Tobago SouthAmerica Ecuador GalapagosIslands WolfandDarwinIslands 3 Colombia MalpeloIsland MalpeloIsland 4 GorgonaIsland GorgonaIsland Fernandode FernandoDeNoronha 3 5 Noronha Europe United UnitedKingdom IsleofMan,Cornwall Kingdom Portugal Portugal CapeSt.Vincent Azores SantaMaria 2 Africa Egypt RedSea RasMuhammad,DaedalusReef,RockyIsland 2 6 Sudan RedSea ElbaReef,PfeifferReef,Sanganeb 4 Tanzania Tanzania PembaIsland Mozambique Mozambique BazarutoArchipelago,PontaD'ouro 4 Madagascar Madagascar NosyBeArchipelago 3 SouthAfrica KwaZulu-Natal SodwanaBay,AliwalShoal,ProteaBanks 3 7 WesternCape DyerIsland,MosselBay,FalseBay Asia Malaysia Malaysia Sabah,BorneoBanks 4 Thailand Thailand KohLantaMarineNationalPark,Phuket 3 Myanmar Myanmar BurmaBanks,MerguiArchipelago 4 India India AndamanIslands 5 Japan Japan OgasawaraIslands 2 Philippines Philippines ApoReef,SuluSea,MaricabanStrait 4 Mauritius Mauritius SharkPit,SharkPoint,SharkPlace 4 Seychelles Seychelles InnerIslands,AldabraAtoll 5 Maldives Maldives FaadhippolhuAtoll,NorthandSouthMaleAtoll 2 5 Australia Australia RaineIsland,OspreyReef,FlindersReef 6 NewSouthWales LordHoweIsland,FishRock,JervisBay 4 Tasmania BassStrait SouthAustralia DangerousReef,NeptuneIslands 43 Province Species Region Country Locality /State AG BL BK HM HR LM LP MK NR RF ST SV SX SM TH SW TG WB WL WH ZB AustraliaCont. Australia WesternAustralia NingalooReef,Kalbarri,RowleyShoals 5 NewZealand NorthIsland Whatuwhiwhi,Tutukaka,HawkesBay 2 SouthIsland Kaikoura SouthPacific PapuaNew PapuaNewGuinea Madang,VitiazStrait,MilneBay 2 8 Islands Guinea Melanesia SolomonIslands RussellIslands,Laulasi,UepiIsland 5 Vanuatu EspirituSantoIsland,NewCaledonia 8 Fiji ManaIsland,KoroIsland,BeqaLagoon 2 6 Micronesia Palau BlueCorner,SharkCity 2 6 MarshallIslands MajuroAtoll,BikiniAtoll 7 Polynesia Marquesas Eiao,Hatutu,Manihi 2 3 BoraBora Tapu,MuriMuri 4 Tahiti Moorea 4 Key:AG=angelshark( Squatinacalifornica ),BL=blueshark( Prionaceglauca ),BK=baskingshark( Cetorhinusmaximus ),HM=hammerheadsharks( Sphyrna spp.),HR=hornshark( Heterodontus spp.), LM=lemonshark( Negaprion spp.),LP=leopardshark( Triakissemifasciatus ),MA=makoshark( Isurusoxyrhinchus ),NR=nursesharks( Ginglymostoma spp.),RF=reefsharks ( Carcharhinus spp.),ST= sandtigers( Odontaspis spp.),SV=sevengillshark( Notorhynchuscepedianus ),SX=sixgillshark( Hexanchusgriseus ),SM=smoothhoundshark( Mustelushenlei) ,SW=swellshark( Cephaloscyllium ventriosum ),TG=tigersharks( Galeocerdocuvieri ),TR=Threshershark( Alopias spp.),WB=wobbegong( Orectolobus spp.),WL=whaleshark( Rhincodontypus ),WH=whiteshark( Carcharodon carcharias ),ZB=zebrashark( Stegostomafasciatum );Numeral=numberofspecieswithintaxon.

44

45