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The Science of Mane-eating lions

The Science of Mane-eating lions

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The Science of Mane-eating lions mentions some interesting info on man-eating lions, as well as other data upon lions in other populations as well.
The Science of Mane-eating lions mentions some interesting info on man-eating lions, as well as other data upon lions in other populations as well.

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Published by: brentlion on Jun 05, 2009
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JournalofEastAfricanNaturalHistory
90:1-40(2001)
THESCIENCEOF'MAN-EATING·'AMONGLIONS
PANTHERALEO
WITHARECONSTRUCTIONOFTHENATURALHISTORYOFTHE'MAN-EATERSOFTSAYO'
JulianC.KerbisPeterhansUniversityCollege,RooseveltUniversity430S.MichiganAve.Chicago,IL,60605-1394,U.S.A.
&
DivisionofMammals,FieldMuseumofNaturalHistoryChicago,IL,60605-2496,U.S.A.jkerbis~frrn]h.orgThomasPatrickGnoskeBirdDivision,ZoologyDepartment,FieldMuseumofNaturalHistory1400S.LakeShoreDrive,Chicago1160605-2496,U.S.A.tgnoske~frrn]h.orgABSTRACTThestoryofthe'Man-eatersofTsavo'hasbeenretoldthroughscript,cinema,andoraltraditioninthe100+yearssincetheirinfamous'reignofterror'.Despitetheirpredictablybroadpopularappeal,thedetailspertainingtothenaturalhistoryoftheselions
Pantheraleo
haveneverbeenreviewed.TheskullsandskinsoftheselionshaveresidedattheFieldMuseumofNaturalHistory(Chicago)forover75years.Ananalysisoftheskulloftheprimaryculpritdisplaysatraumaticinjurythatmayhavelimitedhispredatoryabilityinsubduing'normal'prey.Asampleofhairs,reflectingthedietofbothman-eaters,ispreservedinthebrokenandexposedcavitiesoftheircanines.Variousadditionalcircumstanceslikelycontributedtotheirman-eatinghabit.TheTsavoincidentcloselyfollowedthedebutofrinderpestonthecontinent,whichdevastatedcattleandbuffalo,theprimarypreyoftheTsavolion.TheTsavo'nyika'consistsofadensethornscrubthicketlimitingvisibilityandpassage,representinganidealhabitatforan'ambushpredator'.Finally,historicalreviewoftheliteraturerevealsthat'maneating'wasnotanisolatedincidentatTsavo.Thisbehaviourwaswellestablishedinthevicinityoftherailwaybridgewellbeforetheseinfamouslionsappeared,andcontinuedwellaftertheirdemise,suggestingarecurringopportunity,whichmayhaveevolvedintoalocalbehaviouraltradition.Insum,virtuallyalloftherecognisedpreconditionsforman-eatingoutbreakstooccurwereineffectatTsavointhe1890's.INTRODUCTIONThe'Man-eatersofTsavo'firstgainedinfamyintheBritishpressin1900(TheSpectator,March3)whentheywerereportedtohavecausedatemporaryhalttotheconstructionoftheWeusetheterm'man-eating'byconvention,asattacksarenotlimitedto'men'anddonotalwaysconcludewithconsumption.
 
2
J.C.KerbisPeterhans
&
T.P.Gnoske'Uganda'Railway.ThisrailwaywastorunfromtheIndianOceanportofMombasa,Kenya,totheshoresofLakeVictoria,whichwasthenlocatedinUganda.BritishengineerJ.H.Pattersondocumentedhisattemptstodispatchthetwoadultmalelionswithdetailedjournalentries(Patterson,1898-1899).Theseentriesbecamethebasisforhisfirst-handaccount,
In
theField
(London),firstpublishedattheturnofthecentury,andlaterasthebasisforhisbook,
TheMan-eatersofTsavoandotherEastAfricanAdventures
(Patterson,1907).EveryKenyanschoolboyknowsthestoryoftheTsavolions,firstthroughoraltraditionandlaterthroughthispopularreference.TwoHollywoodfilmshavefurtherpopularisedthestory:
BwanaDevil
(1953)and
TheGhostandtheDarkness
(1997).Despitethepopularappealofthisincident,nodetailsconcerningthelionsthemselves,orthecircumstancessurroundingthisnotoriousman-eatingoutbreak,werepublishedbetweenPatterson's1907accountandthebriefreviewbyKerbisPeterhans
etal.
(1998).Afterreviewinghistoricalaccounts,recentliterature,theoriginaljournalsofLH.Patterson,GameDepartmentrecords,andtheskullsandskinsofthe'Man-eaters'themselves,wehaveidentifiedthesecircumstances.Thispaperaddressessomeoftheconditionsthatleadtoman-eatingingeneralandhowtheserelatetotheincidentsatTsavoover100yearsago.Severalcircumstanceshaveoftenbeenassociatedwithpantherid(bigcat)attacksonhumans.Sickly,injured,agedoremaciatedpantherids,whichareunabletosecuretheirnormalprey,canoftenbecomeman-eaters(Corbett,1944,1948,1954).Predatorsmayalsofocusonabnormallybehavingprey,withcertainhumanbehavioursprobablyqualifying.Amongsocialcarnivores
(e.g.
lions),thefavouringofparticularpreyspecies
(e.g.
livestockorhumans)canbepassedfromonegenerationofpredatorstothenext,withthepotentialtobecomealocalsocialtradition(Swayne,1895;Taylor,1959;Rushby,1965).HistoricalrecordsfromTsavosuggestthatpredationuponhumanswasalong-standingphenomenon.HumanswereattackedandkilledbylionsintheTsavoareawellbeforeconstructionoftherailwaybegan(1886,documentedinJackson,1894)andcontinueatpresent,over100yearsaftertheinfamouspairwasdispatched(KenyaWildlifeService,1994-1998).Anotherfactor,anecdotallymentionedintheliterature,istheabsence,depletion,orremovaloftypicalprey,causingthepredatorstoseekalternativefoodsourcessuchaslivestock.Thisbringslionsintocontactwithhumans,sometimesresultingintheinclusionofhumansinlions'diets.Lionsalsodevelopatasteforhumansafterbeing'provisioned'withdeadones.Environmentalvariablescanbeimportantsinceheavycoverisessentialfor'ambushpredators'tosuccessfullystalktheirprey(Schaller,1972;Funston
etai,
2001).Seasonalfactorsarealsothoughttobeimportantasgamedispersesduringtherains,makingthemmoredifficulttolocateandsecure(Jackson,1894).Inlate19thcenturyTsavo,allofthesefactorswereinplayconcurrentlyandallmayhaveplayedaroleinthedevelopmentoftheworld'smostrenownedman-eatingoutbreak.Whatfollowstherefore,isacriticalreviewofthesedifferentscenariosandthelikelyroleeachofthemmayhaveplayedintheeventsatTsavoover100yearsago.NOTESONTHENATURALmSTORYOFTHE'MAN-EATERSOFTSAVO'UnliketheTsavooftodaywithlargetractsofopenexpanse,theTsavoofthe1890'swascomposedofanearlyimpenetrable,thornthicketknownas'nyika'.Thequestforivoryduringthe19thcenturyhadeliminatedelephants
(Loxodontaafricana)
frommuchofeasternKenya,includingmostofTsavo(Thorbahn,1979).ThisiswhymanyoftheporterstransportingivorythroughTsavoatthetime,originatedfarup-country,sometimesevenfrom
 
TheScienceof'Man-Eating'amonglions
PantheraLeo
3Uganda(Patterson,1907,Preston,n.d.).Elephantsareakeystonespeciesandhaveamajorimpactonthevegetationandthelargemammalcommunity.TheeliminationofelephantsfromTsavocausedaproliferationofdensewoodyandthornyundergrowthandeliminatedherdsofgrazingungulatesfromthevicinity.Preston(n.d.)describesthisthickhabitatindetail.Browsers,includingdik-dik
Madoqua
sp.andblackrhino
Dicerosbicomis
increased.AreviewofPatterson'sfieldjournals(1898-1899)quantifiestheanimalspeciesheencountered(table1).Therewerenowildebeest
Connochaetestaurinus.
Hartebeest.(hartebeest/hirola,
Alcelaphusbuselaphus/hunteri)
andzebra
Equusburchelli/grevyi
werearoundbutnotcommon.Pattersonneveroncereferstobuffalo
Synceruscaffer
orcattle
Bostaurus
ssp.inhisjournal.Thiscanbetracedtothearrivalofrinderpestonthecontinentin1891,whichhadadevastatingimpactonthebovinepopulations(Lugard,1893;Mettam,1937).Bythe1960's,theTsavoelephantpopulationhadrecoveredandTsavobecameknownasoneofthelargestelephantsanctuariesinEastAfrica(Leuthold
&
Leuthold,1976).Therefore,thetwolargemammalspecies(elephantandbuffalo)comprisingmostofthemammalianbiomassinTsavotoday
(ibid.),
werevirtuallyabsentinthe1890's.Insum,by1898,Tsavohadalreadybeenseverelyimpactedbyhumans,resultinginvastanthropogenicdifferencesbetweentheTsavoenvironmentofthe1890'sandthatoftoday.The'Man-eatersofTsavo'themselvesconsistoftwoadultmalelionsthatarevirtuallydevoidofmanesintheconventionalsense.Theydisplayaslightsagittalcrestofhairthatcanbetypicalofsomemanelesslions.Thesecondlionkilled(FMNH23969,figure1)hadslightcheektuftsaswellasdarkenedchestpatcheswhilethefirstlionwaswithouttheseadornments.TheywerephotographedbyColonelJ.R.Patterson(1907)aftertheyweredispatchedin1898.Exceptfortheseverelybrokencanineandremodeledmandibleandcraniumoftheprimaryculprit(FMNH23970;figure2),theman-eatersofTsavowereadultmalelionsintheirprime(6.5-8.5yearsofage).Theirsquamoso-parietalsuturesareobliteratedmakingthematleast6.5yearsold(Smuts
etal.,
1978),buttheirmaxillopremaxillarysuturesarenotevenpartiallyclosed,makingthemunder9yearsold.Theirteethalsobetraythisagerangeastheyareyellowingandthereisvisiblewearonthecanine,incisor,
p3
andP4
(>
5-6yearsofage,
ibid.).
Althoughtheseindividualsmayhavebeensiblingsduetotheirassociationandsimilarsizeandmorphology,FMNH23969displaysmoreapicalwearthanFMNH23970.Wedoubtthattheirdifferenceinageextendsbeyondoneortwoyears.BothanimalswerehugeindividualsasPatterson'spublishedphotos(1907)demonstrate.Pattersonusesthefollowingindescribingthefirstlion:"enormousbrute","powerfulbeastineveryway",(Patterson,1898-1899).FMNH23970was2.95mlong(nosetotail)and1.12minheightattheshoulderwhileFMNH23969was2.90mlongand1.22minheightattheshoulder
(ibid.).
EastAfricanmalelionsaverage0.96mheightatshoulderand2.67minlengthaccordingtoMeinertzhagen(1938)whocollectedintheAthi-KapitiPlainsandintheSerengeti.Guggisberg(1975)providesanadditionalmeasurement:0.91minheightattheshoulderand2.74mlong.Patterson(1907)claimedthattheman-eaterswereprime-aged,healthymalesandwassocitedbysubsequentauthors
(e.g.
Selous,1908;Akeley,1923;Bradley,1926;Guggisberg,1961;Cloudsley-Thompson,1967).Wecanonlyassumethatthisreferredtotheconditionoftheirlimbsandtorsoandthathedidnotinspecttheirdentition.ResidinginthecollectionsoftheFieldMuseumofNaturalHistory(FMNH)for75years,thetwoTsavolionskullshadnotbeendifferentiatedfromoneanotherinanyway.Oneofus(TPG)rediscoveredtheseskullsinthecollectionsoftheFieldMuseuminthe1980'sandsubsequentlydeducedwhichwasthefirstman-eatershotbyreferringtoPatterson'sdiscussionofthedeathsofeach

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