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ScavengingbychimpanzeesatNgogoandtherelevanceofchimpanzeescavengingtoearlyhomininbehavioralecologyDavidP.
WattsDept.
ofAnthropology,YaleUniversity,P.
O.
Box208277,NewHaven,CT06511-8277,USAReceived28February2007;accepted3July2007AbstractChimpanzeesregularlyhuntavarietyofpreyspecies.
However,theyrarelyscavenge,whichdistinguisheschimpanzeecarnivoryfromthatofsomemodernhunter-gatherersand,presumably,atleastsomePlio-Pleistocenehominins.
Iuseobservationsmadeoveran11-yearperiodtodocumentallknownopportunitiesforscavengingencounteredbychimpanzeesatNgogo,KibaleNationalPark,Uganda,anddescribeallcasesofscavenging.
Ialsoreviewdataonscavengingfromotherchimpanzeeresearchsites.
ChimpanzeesatNgogoencounteredscavengingoppor-tunitiesonlyaboutonceper100daysandatemeatfromscavengedcarcassesonlyfourtimes.
Scavengingopportunitiesarealsorareatothersites,evenwhereleopardsarepresent(Mahale,Ta,Gombe),andscavengingofleopardkillsisknownonlyfromMahale.
Feedingonpreythatchimpanzeeshadhuntedbutthenabandonedisthemostcommonformofscavengingreportedacrossstudysites.
Forexample,severalindivid-ualsatNgogoatemeatfromapartiallyconsumedredcolobuscarcassabandonedafterahuntthepreviousday.
SuchbehaviorprobablywasnotcommonamongOldowanhominins.
Ngogodataandthosefromothersitesalsoshowthatchimpanzeessometimeseatmeatfromcarcassesofpreythattheydidnotseekilledandthatwerenotkilledbychimpanzees,andthatscavengingallowsaccesstocarcasseslargerthanthoseofanypreyitems.
However,chimpanzeesignorerelativelymanyopportunitiestoobtainmeatfromsuchcarcasses.
Scavengingmayberarebecausefreshcarcassesarerare,becausetheriskofbacterialinfectionsandzoonosesishigh,andbecausechimpanzeesmaynotrecognizecertainspe-ciesaspotentialpreyorcertainsizeclassesofpreyspeciesasfoodsources.
Itsminimalnutritionalimportance,alongwiththeabsenceoftech-nologytofacilitateconfrontationalscavengingandrapidcarcassprocessing,apparentlydistinguisheschimpanzeeforagingstrategiesfromthoseofatleastsomeOldowanhominins.
2007ElsevierLtd.
Allrightsreserved.
Keywords:Chimpanzees;Meateating;Oldowanhominins;ScavengingIntroductionWildchimpanzeesregularlyhuntandconsumeavarietyofvertebrateprey,andtheyeatmoremeatthandomostorallothernonhumanprimates.
Detaileddataonhuntingandmeatconsumptionhavebeencollectedatmultiplesites,espe-ciallyatGombe(Goodall,1986;Stanford,1998),Ta(BoeschandBoesch,1989;BoeschandBoesch-Achermann,2000),Mahale(Hosakaetal.
,2001),andNgogo(MitaniandWatts,1999,2001;WattsandMitani,2002).
Chimpanzeessometimesopportunisticallypursuesolitaryprey,butmoreoftenconductgrouphuntsofgregariouspreyinwhichtheymaydeploycom-plex,cooperativeprey-capturetactics(BoeschandBoesch,1989;BoeschandBoesch-Achermann,2000).
Theyalsocom-monlysharemeataftersuccessfulhunts.
Comparativedataonchimpanzeehuntingandmeatconsumptionarevaluableforinvestigationsoftheimportanceofhuntingandmeateatinginhumanevolution(e.
g.
,Moore,1996;Stanford,1996,2001;Kaplanetal.
,2001)andtheevolutionaryecologyoffoodsharingbyhumans(e.
g.
,Winterhalder,2001).
Incontrast,reportsofscavengingbychimpanzeesarerare.
Inthispaper,IsummarizepublisheddataonscavengingbychimpanzeesandpresentnewdatafromNgogo,KibaleNa-tionalPark,Uganda.
IdiscussthesedatawithreferencetothedebateovertherelativeimportanceofhuntingandE-mailaddress:david.
watts@yale.
edu0047-2484/$-seefrontmatter2007ElsevierLtd.
Allrightsreserved.
doi:10.
1016/j.
jhevol.
2007.
07.
008Availableonlineatwww.
sciencedirect.
comJournalofHumanEvolution54(2008)125e133scavenginginthesubsistencestrategiesofPlio-Pleistocenehominins.
ChimpanzeesatNgogohuntfrequentlyandareun-usuallysuccessfulpredatorsofredcolobusmonkeys(MitaniandWatts,1999,2001;WattsandMitani,2002),buttheyig-noremostoftheirrareopportunitiestoscavenge,althoughtheyhavebeenobservedtoeatpartsoffourcarcasses.
Thisseemstypicalofchimpanzeesinforesthabitats,andthecon-tributionofscavengingtochimpanzeesubsistenceisminimal.
ScavengingbymodernhumansandPlio-PleistocenehomininsThesubsistencestrategiesofsomemodernhumanhunter-gatherersincludescavenging.
Thebest-studiedexampleistheHadza,whomonitorlargecarnivoresandvulturestolearnofscavengingopportunitiesthattheyexploiteitherpassivelyorbyconfrontingthecarnivores,includinglions,andstealingtheirkills(O'Connelletal.
,1988a;Bunn,2001).
Ofallthemedium-to-large-sizedanimalcarcassesthatagroupofHadzaforagersacquiredduring14monthsin1985e1986,20%werescavenged(O'Connelletal.
,1988a).
Confrontationalscaveng-ingledtoacquisitionofcarcassesthatwereover75%completeinveof11scavengingcasesandoccasionallypro-videdlargeamountsofmeat.
However,seasonalvariationincarcass-encounterratesandvariationinthesizeandcomplete-nessofcarcassesledtohighvarianceinreturnsfromscaveng-ingandmadescavengedanimaltissueonlya''windfallresource.
''Inaddition,passivescavengingyieldedlittlemeat(O'Connelletal.
,1988a).
Extensivedebatesurroundsthequestionsof(1)howimpor-tantscavengingwas,comparedtohunting,inPlio-Pleistocenehomininsubsistencestrategiesand(2)wheninthecourseofhumanevolutionmeatbecameacriticaldietarycomponent(re-viewedbyDominguez-Rodrigo,2002;O'Connelletal.
,2002;Dominguez-RodrigoandPickering,2003;Plummer,2004).
Thisdebatelargelyinvolvescontrastinginterpretationsofre-sultsfromanalysesoffaunalassemblagesfromOldowansitesthatshowhominin-inducedmodication.
Italsoinvolvesdis-agreementabouttherelativecontributionsoffoodsofanimalorplantorigintothedietsofmodernhumanhunter-gatherersand,concomitantly,(1)therelativeimportanceofmen'sandwomen'scontributionstohunter-gatherersubsistence;(2)theroleofhuntingandmeatsharinginmalereproductivestrategies;and(3)therelevanceofethnographicexamplestore-constructionsofearlyhomininbehavioralecology.
Iwillonlybrieyreviewsomeaspectsofthisdebatetowhichchimpanzeedataarepotentiallyrelevant.
IfanyOldowanhomininsfre-quentlyscavenged,andscavengingcontributedsubstantiallytotheirdiet,thenthiswouldhavebeenamajordifferenceinbe-havioralecologyfromthatofextantchimpanzees.
Inaddition,ifscavenginghadonlyminornutritionalimportance,butcon-frontationalscavengingofcarnivorekillswasimportantasamalematingtactic(O'Connelletal.
,2002),thenthiswouldraisetheissueofwhether,andifso,how,meatsharingentersintothereproductivestrategiesofmalechimpanzees.
DeterminingthetimingofaccesstocarcassesandhowmuchmeattheycouldhaveprovidediscrucialtoassessinghowOldowanhomininsassembledmodiedanimalbones.
Manykindsofdataarerelevanttosuchdeterminations,in-cludingthoseonthefrequencyandlocationofstone-toolcutmarksandcarnivoretoothmarks,onbone-fracturepatterns,onskeletal-partproles,andonhowextantcarnivoresandcontemporaryhumanforagers(e.
g.
,theHadza:O'Connelletal.
,1988b;Bunn,2001)modifycarcasses.
Extensiveanaly-sesofungulateremainsfromseveralBedIsites(especiallyFLKIZinjanthropus)atOlduvaiGorge,Tanzania,mostofwhicharefrompreyofsizeclasses2or3(adultbodyweight40e300kg)andwhichdatetoabout1.
75millionyearsago(Ma),haveledtocontrastinginterpretations:(1)homininshadonlylateaccesstocarcassesandprobablyonlyextractedmarrowfrombonesmostlydeeshedrstbycarnivores(Blu-menschine,1991),or(2)theyregularlyhadearlyaccesstocar-casses,consistentwitheitherhuntingortheftofcarcassesfromlargecarnivores(BunnandKroll,1998;Dominguez-Rodrigo,1997,1999;Bunn,2001;O'Connelletal.
,2002;Bunn,2006).
DatafromseveralotherPliocenesitesareconsistentwithearlyaccesstocarcasses.
Inparticular,highfrequenciesofcutmarksonlong-bonemidshaftsandothermeatyskeletalelements,combinedwithlowfrequenciesofcar-nivoretoothmarksontheseelements,pointtoearlyaccessatOlduvaisiteFxJj50(Dominguez-Rodrigo,2002),Peninj(Dominguez-Rodrigo,2002),andperhapsatGona(Domi-nguez-Rodrigoetal.
,2005).
InarecentreanalysisofdatafromFLK-Zinjanthropusandcritiqueofearlieranalyses,Dom-inguez-RodrigoandBarba(2006)concludedthathomininsalsohadearlyaccesstothecarcassesofmedium-sizedbovidsaccu-mulatedatthissite.
Earlycarcassaccessdidnotnecessarilyentailhunting.
O'Connelletal.
(2002)agreedthatbone-modicationdataandactualisticpatternsshowearlyaccesstomanycarcasses,butcontendedthatlackofprojectiletechnologypreventedPlio-Pleistocenehomininsfromhuntinglargegame,andthattheygainedearlyaccessbyaggressivelyconfrontingpredatorsatrelativelyfreshkills.
Likewise,Stiner(2002:6)arguedthat,whilesomehuntingofsmallerpreyprobablyoccurred,largemammals''wouldhavebeendifculttokillwithOldowanorearlyAcheuleantechnology,''althoughshealsonotedthattheinitialoccupationofpartsofEurasiabyhomininsthatpossessedsuchtechnologydependedonforagingstrategiesthatincludedcarnivory.
However,Dominguez-Rodrigoetal.
's(2001)plant-residueanalysisofAcheuleantoolsfromPeninjprovidedevi-denceforwoodworkingat1.
5Ma,fromwhichtheyconcludedthatHomoergasterhadtheabilitytoproducespearsthatcouldhavebeenusedforhunting.
O'Connelletal.
(2002)alsoarguedthathighcarnivoretooth-markcountsonbonesfromOldowansitesindicateaggressivescavengingofcarcassesalreadyde-eshedtovaryingdegrees,nothunting.
Dominguez-Rodrigo(2002,2003;cf.
Dominguez-RodrigoandPickering,2003)counteredbynotingthatthehighfrequencyofcutmarksonboneswherecarnivoresstartdeeshing(e.
g.
,upper-limblongbones,pelves),andespeciallythehighfrequencyofcutmarksonupper-limblong-boneshafts,resemblesthepatternthatre-sultswhenhumansinitiatethedeeshing,ashappenswhentheyhavehuntedtheanimals.
126D.
P.
Watts/JournalofHumanEvolution54(2008)125e133SeveralresearchershavearguedthatmeatobtainedfromconfrontationalscavengingwasnotamajorcomponentofOldowanhominindiets,foravarietyofreasons.
Forexample,Blumenschine(1991)arguedthattherisksoftryingtostealfreshcarcassesfromlargecarnivoreswouldhavebeenex-tremelyhigh(withtheexceptionofcachedleopardkills)andthattheoccurrenceofscavengingopportunitieswouldhavevariedgreatlyonaseasonalbasis.
Othershavearguedthatconfrontationalscavengingwouldhaveoccurredonlyifthehomininshadconsiderablenumericalsuperiorityovertheircarnivorecompetitorsand/oriftheycouldreducetheriskssuf-cientlybyusingweapons(Shipman,1986;O'Connelletal.
,1988a,b;VanValkenburgh,2001).
BasedontheirstudyofHadzascavenging,Hawkesetal.
(1988a)assertedthathighvarianceinthetimingofopportunitiesandinpossiblereturnswouldhavemadeconfrontationalscavengingatbestasourceofoccasionalwindfallsforOldowanhominins,notaregularactivity.
AstheirexampleofHadzascavengingofanearlyin-tactgiraffecarcassindicated,however,returnsmightsome-timeshavebeenextremelyhigh.
AgainrelyingonHadzadata,O'Connelletal.
(2002)arguedthatconfrontationalscav-engingbymalesservedmainlyascostlysignalingthatgavefemalesaccurateinformationaboutmatequality,ratherthanasanimportantsourceofnutrients.
Intheirview,meatwasanunreliablefoodsourceanditsdietaryimportancesecondarytothatofplantfoods,particularlyundergroundstorageorgans.
However,othershavecautionedagainsttakingtheHadzaeitherasgeneralrepresentativesofmodernhunter-gatherersorasanalogiesforOldowanhominins(e.
g.
,Kaplanetal.
,2001;Dominguez-Rodrigo,2002,personalcommunication).
Areviewoftheethnographicrecordsupportstheclaimthatmostmodernhunter-gatherersgainedmostoftheircaloriesfrommeat,andthusfromhunting,amostlymaleactivity(Kaplanetal.
,2001).
Dominguez-Rodrigo(2002;c.
f.
Dominguez-RodrigoandPickering,2003)attributedmajorim-portancetomeatinOldowanhomininsubsistence,butarguedthatmostofitcamefromhunting,notconfrontationalscavenging.
Plummer's(2004)preliminaryanalysisofprey-sizedistributionandevidenceforbutcheryandcarnivoreac-tivityatthe2-million-year-oldsiteofKanjerahelpstoshiftthedebateawayfromalongstandingfocusonlargegame.
Amongotherlinesofevidence,highfrequenciesofstone-toolcutmarks,lowfrequenciesofcarnivoretoothmarksonlimb-boneshafts,andhighrepresentationofjuveniles(usuallycompletelydestroyedbycarnivores)indicatethathomininsregularlyhuntedsmall(10a17!
118Scavengingcases10b7c7b4Casesfromearlierredcolobushunts5e9d30e1dScavengenonpreyspeciesNoNoYes(bushbuck)NoExtendpreysizerangeNoNoYesYesIgnorelargepreyYes(bushpig,bushbuck)(duiker)eYes(bushpig)YesStealkillsbyotherpredatorsBaboonCrownedeagleLeopardCrownedeagleConfrontationalcarcasstheftYesYesNoNoFeedoncarcasseswhenpredationunseenNoNoYesYesSources:Gombe:Goodall(1986),Mulleretal.
(1995),W.
Wallauer(personalcommunication);Ta:BoeschandBoesch-Achermann(2000);Mahale:Hasegawaetal.
(1983),Nishida(1994);Ngogo:thisstudy.
aNotincludingcarcasstheftsfrombaboonsorencounterswithbaboonseatingmeat(Goodall,1986;numberofcasesnotreported).
bChimpanzeesobtainedonecarcassfromhumansatGombe(Goodall,1986)andtwofromhumansatMahale(Hasegawaetal.
,1983).
cDoesnotincludethreecasesinwhichchimpanzeesstolelivingpreyfromcrownedeagles.
dFivedeniteandfourprobableGombecases;informationnotprovidedforonecaseatMahale.
eChimpanzeesatTaignoreduikerkilledbyleopards,butalsodonothuntduiker.
127D.
P.
Watts/JournalofHumanEvolution54(2008)125e133casesofscavenginghavebeenobservedatGombesinceGoodall's(1986)review(W.
Wallauer,personalcommunica-tion).
Hasegawaetal.
(1983)reportedfourcasesofscaveng-ingatMahale,twoofadultbushback(Tragelephusscriptus)probablykilledbyleopards(onenearlyintact,thesecondfreshbutmostlyeaten)andtwoofblueduiker(Cephalophuscalli-pyga)carcassesthatresearchershadhandled.
Nishida(1994)laterreportedthreemorecases:oneblueduiker,onered-tailedmonkey(Cercopithecusascanius),andoneredcolobus(Pilio-colobustephrosceles).
Basedonobservationsovernearlytwodecades,BoeschandBoesch-Achermann(2000)reportedonlysevencasesatTa,includingthreeoffeedingonremainsofmonkeyskilledbychimpanzeesoneortwodayspreviouslyandfouroffeedingonmonkeyskilledbycrownedeagles(Ste-phanoaetuscoronatus).
Chimpanzeesalsostolewoundedredcolobusfromeaglesthreetimesandencountered,butignored,tenfreshcarcassesofseveralmammalianspecies,althoughonlyoneofthese(sootymangabeys,Cercocebusatys)wasaknownpreyspecies.
Reynolds(2005)reportedonecasefromBudongo,inwhichchimpanzeesfedonthecarcassofaninfantbluemonkey(Cercopithecusmitis)thathadbeenkilledandpartiallyconsumedbyanadultmalebluemonkey;chimpanzeesoccasionallypreyonbluemonkeysatthissite.
ScavengingbychimpanzeesatNgogoMethods:studysiteandsubjectsNgogoisinthecenterofKibaleNationalPark,inwesternUganda.
Thestudyarea,atanaltitudeofabout1400m,receives1700e1800mmofrainannually,mostlyfromMarchthroughMayandmid-AugustthroughDecember.
Itcoversabout30km2ofmixed,mature,andregeneratingforestthatistransi-tionalbetweenlowlandandmontaneevergreenforestandincludesother,minor,vegetationtypes(Butynski,1990;Struhsaker,1997;Lwangaetal.
,2003).
Threesmall-to-medium-sizedungulatesarecommon(Struhsaker,1997):redduiker(Cephalophusmonticola),blueduiker,andbushpig(Potomochoerusporcus).
Bushbuckarealsopresent,butarelesscommon.
Sevenotherdiurnalprimatespeciesarepresent,withred-tailedmonkeysandgrey-cheekedmangabeys(Lophocebusalbigena)beingthemostcommon(Mitanietal.
,2000).
Thechimpanzeecommunityhasbeencontinuouslyob-servedsincemid-1995andisthelargestknown;its140e150membershaveincluded22e28adultmalesand14e16adolescentmales(Watts,1998;MitaniandWatts,2005).
Thechimpanzeespreyonatleast11vertebratespecies,in-cludingsixdiurnalprimatesandthethreecommonungulates(althoughtheydonothuntadultbushpigsoradultreddui-ker).
Thepredationdatabaseincludesover650killsduringmorethan270hunts(WattsandMitani,2002,unpublisheddata).
Redcolobusmonkeysarebyfarthemostcommonprey;about80%ofredcolobushuntsresultinkills,andchimpanzeesmakemultiplekillsofredcolobusinmosthunts.
IobservedchimpanzeesatNgogoforover10,000hoursbe-tween1995and2006.
Irecordedallhuntsandpreycaptures,collecteddataonmeatsharing,andnotedallencounterswithfreshprey-speciescarcassesnotkilledbychimpanzeesandallcasesofscavenging.
ResultsChimpanzeesencounteredonlyeightrecentlydead,nonde-composedcarcassesofpreyspeciesandfoundoneindividualofapreyspeciesdyingfromunknowncauses.
Thus,theen-counterratewithscavengeablecarcasseswasonlyonceperca.
98days,assuminga13-houractivityperiod.
Thisincludestwocasesinwhichadultmalebaboonswereeatingredduikerthattheyhadjustkilled;thechimpanzeesshowednointerestintryingtostealthesecarcasses.
Intwoothercases,chimpan-zeesencountereddeadadultredduikerthattheydidnotcon-sume.
Intherstofthesecases,threemalechimpanzeesignoredthecarcass,andinthesecond,anadultfemalewithaninfantandanadolescentfemalebrieyinspectedthecar-cass,butdidnottouchit.
Thefourscavengingepisodes,andoneinteractionbetweenanadultmalechimpanzeeandadyingblack-and-whitecolobusmonkey(Colobusguereza),aredetailedbelow.
Case1:adultfemaleredduikerAt0920honSeptember26,2002,adolescentmalesRAandWAandadolescentfemaleBIfoundadeadadultfemaleredduiker.
Itsabdomenhadanarrow6-cmopeningandsomeviscerahadbeenremoved;nootherwoundswerevisibleandthecarcasswasotherwiseintact.
Thelimbswereslightlyex-ible,butmanyieswereswarmingaroundthecarcass.
WAandBIinspecteditcloselyandsniffedit,butdidnottouchit.
RAapproachedit,sniffedandvisuallyinspecteditandpokedatitsventrum.
Hepulledandtwistedoneoftheduiker'slegswithbotharmsforfourminutesasiftryingtodisarticu-lateit,pokedattheopeningintheabdomenagainandsniffedhisnger,thendraggedthecarcass3mandpulledandtwistedthelegforanotherveminutes.
Finallyhebitintothelegsev-eraltimes,piercingtheskinjustabovethekneeandremovingasmallamountofmeat,whichheate;hedidnotbiteintothemeatierhipregion.
Hesatnexttothecarcassforseveralmoreminutesandthenleftthesite.
WAfollowedhim,butBIthenbrieypulledandtwistedoneleg,bitintoit,andremovedasmallamountofmeat,whichsheate.
Shethenfollowedthemales.
SofarasIwasaware,nootherchimpanzeesencounteredthecarcass.
Case2:adultfemaleredcolobusAt1645honDecember10,2002,15adultmalechimpan-zees,veadolescentmales,12adultfemales(sevenwithin-fants),twoadolescentfemales,andninejuvenilesencounteredaredcolobusgroup.
Theystartedtohuntthemonkeysat1710handmadevekills(oneadultfemale,twosubadults,andtwoinfants)betweenthenand1755h.
Onemoreadult128D.
P.
Watts/JournalofHumanEvolution54(2008)125e133malechimpanzeeandoneadolescentmalearrivedduringthesubsequentmeat-eatingsession.
At1820h,chimpanzeescreamswereaudiblefromthenorth;thecallersbelongedtoaneighboringcommunity.
TheNgogomalesbecamehighlyexcitedandrushedinthatdirec-tion.
Atleasttwomaleswerecarryingmeat(thetwolargestcarcasses),butdroppeditastheyapproachedthecallers.
At1842h,themalesstoppedandlistenedonthesouthernedgeofasmallvalley;chimpanzeescalledfromthenorthsideofthevalleyat1845h.
TheNgogomalesexcitedlybutsilentlyapproached,andat1858htheychargedtheotherchimpan-zees,whoscreamedandednorth.
Malescontinuedtodisplayuntil1915h,thenmovedsouthandnestedintheareaofthehunt.
Onthenextmorning,11oftheadultmales,threeadoles-centmales,oneorphanedjuvenilemale,andseveralfemales,somewithdependentoffspring,returnedtothesouthernedgeofthevalleywherethemaleshadattackedtheneighborsthepreviousevening.
At1303h,allthemalesplusoneoldfemalestartedapatroltothenorthandeast,duringwhichtheytrav-eledseveralkilometerswithoutencounteringneighbors.
At1205h,beforethepatrolstarted,adultmaleORfoundmostofanadultredcolobuscarcass,presumablythefemalekilledthepreviousafternoonbyHA,ontheground.
Hepickeditup,carriedit20m,thenclimbedatreeandstartedtoeatmeat.
Meanwhile,adultmaleAYfoundpartofalimb,appar-entlyfromthesamemonkey,andstartedtoeatmeat.
ORsharedmeatwithtwoadolescentmales.
One(DX)brieyre-trievedthecarcassandatemoremeatwhenORdroppeditat1240tofollowothermalesacrossthevalley.
Thesefourmalestreatedthecarcasslikeafreshkill:nonehesitatedtoeatmeat,althoughtheydidnoteatallofit.
Nootherchimpanzeesshowedinterestinobtainingmeat.
Case3:subadultred-tailedmonkeyOnthemorningofJuly12,2005,14adultmales,6adoles-centmales,anestrousfemaleandheradolescentandjuveniledaughters,andatleastsevenotheradultfemaleswithveju-venilesandfourinfantswereeatingChrysophyllumwarwickiifruitinseveraltrees.
At1310h,youngadultmaleGTZfoundadeadsubadultmalered-tailedmonkeyonthegroundbeneathoneofthesetrees.
Itappearedfreshditslimbswereexibledandhadnoobviouswounds(althoughthoseinictedbycrownedeaglesmaybevisibleonlyoncloseinspection;T.
Struhsaker,personalcommunication).
GTZgraspeditstailanddraggedit10masadolescentmaleMUwatched,thenstoppedandbitintooneofitshipjoints.
Afterseveralminutesofbitingandpullingatthejoint,hetoreoffonelegandthetail,carriedtheseupatree,andstartedtoeatmeat.
Noonejoinedhimorbeggedformeat.
AdultmaleLO,whoarrivedwhileGTZwasdisarticulatingthelegandtail,draggedtherestofthecarcass20malongtheground,thensatandlookedatitfor3min,butatenomeat.
Nootherchimpanzeesinves-tigatedthecarcass.
GTZwasstilleatingmeatat1400h,whenfouradultmalesandseveralfemalesandtheirdependentoffspringmovedsouthandencounteredaredcolobusgroupabout1kmfromthescavengingsite.
Onemaleclimbedtowardtheredcolobus,butthemonkeysed.
GTZ,stillcarryingpartofthered-tailedmonkeycarcass,arrivedwithseveralothermalesat1510h.
Heclimbedabout12mupaPseudospondiasmicrocarpatreeandresumedeatingmeat.
Anadolescentfemalejoinedhimandbeggedunsuccessfullyformeat.
At1605,thechimpanzeeshuntedblack-and-whitecolobusinthesameareaandadultmaleMOkilledajuvenile,halfofwhichanotheradultmalestole.
GTZdidnotparticipateinthehuntorjoinbegging/meat-sharingclustersthatformedaroundthetwomeatpossessors.
Hedroppedtheremainsofthered-tailedmonkeysometimebetweenthenand1700h.
Case4:subadultredduikerAt1215honApril4,2006,alargepartyofchimpanzeesencounteredthecarcassofasubadultredduikerlyingonthegroundinaswampyvalley.
Theduikerappearedtobefreshlydeadandmighthavebeenkilledbyacrownedeagle.
Sixadultmalesplusseveralyoungadolescentmalesandmanyfemales,juveniles,andinfantscrowdedaroundthecar-cassandlookedatit.
Noneoftheadultstoucheditatrst,butseveraljuvenilestouched,sniffed,andhitit.
JuvenilemaleBNstartedtodragitaway;thispromptedalphamaleBTtochargethroughthemiddleoftheassembledgroup.
Mostofthechim-panzeesed,andmanyothersalsochargedatorotherwiseen-gagedinaggressiveinteractions.
Duringthismelee,BNletgoofthecarcassandayoungadolescentfemaledraggeditintothickvegetation,butthenalsoletgoofit.
Atsomepoint,adultmalePAseizedthecarcassandcarrieditupaPseudospondiasmacrocarpatree,wherehestartedtoeatit.
Sixotheradultmales,oneadolescentmale,andonejuvenilemalefollowedandstartedtobegformeat.
PAwasreluctanttoshareandman-agedtopreventallbutalphamaleBTfromobtainingmeatwhilehefedontheduikerforoveranhour.
BTleftduringthistime,buttheothermalesbecameincreasinglyagitated,andnally,adultLOchargedatPAandprecipitatedanothermelee,duringwhichhestolethecarcassfromPA.
AdultmaleMORthenstoleitfromhim.
MORsharedmeatwiththreeotheradultmalesandanadolescentfemale,andrefusedbeggingattemptsbyafourthadultmale.
Oneotheradultmaleandayoungadolescentmaleobtainedscrapsofmeatfromtheground.
Theentiremeat-eatingsessionlastedalmostfourhoursandthechimpanzeesconsumedtheentirecarcass.
Case5:subadultmaleblack-and-whitecolobusAt1600honJuly1,2002,alphamaleBTlefteightotheradultmales,twoadolescentmales,andonefemalewithajuve-nileandinfanttoforagealone.
At1610h,heheardalarmcallsfromagroupofblack-and-whitecolobusmonkeys;helookedtowardsthemonkeys,thenrushedoverandsatbelowthem.
Suddenlyhespottedalargesubadultmalemonkeyontheground,alivebutnotmoving.
Itwasnotobviouslywounded,andmighthavefallenbecauseofanaccident,becauseofag-gression,becauseitwasill,orbecauseofacrownedeagle129D.
P.
Watts/JournalofHumanEvolution54(2008)125e133attack.
BTdraggedthemonkey30mbyitstail,thrashingitagainstthegroundseveraltimes;itresistedineffectivelybygrabbingatplantstems.
Afterstoppingtositandlookatit(withoutreleasinghisgrip),hedraggeditanother10m,thrasheditagainstthegroundagain,andcarrieditupaPseu-dospondiasmicrocarpatree,whereheplaceditonalargeboughandsatnexttoit.
Hehitthemonkeyagainsttheboughseveraltimes,sniffeditandinspecteditvisually,andputhismouthonitshead,butdidnotbiteit.
Hethencarriedittoasecondbough,putitdown,andsatlookingcloselyatit.
Themonkeywasnolongermovingandappeareddead.
Tenminutesafterherstclimbedthetree,BTdescendedandmovedoffwithouthavingeatenanymeat;heleftthemonkeyinthetree.
DiscussionTheseobservationsaddtothesmallbodyofdatashowingthatwildchimpanzeesinlowlandevergreenmoistforests(Ta),mid-altitudeevergreenmoistforest(Kibale,Budongo),anddryer,moreseasonalforests(Mahale)andwoodland(Gombe)scavengevariousvertebratespeciesonwhichtheyalsoprey.
However,scavengingisrare,especiallycomparedtoratesofpredationonsomeofthesamespecies,andchim-panzeesdeclinealargeproportion,andmaybethemajority,ofscavengingopportunities(Table1).
Oneobviousreasonfortherarityofscavengingistherarityofopportunities.
Theestimateofoneencounterper98daysatNgogoisconsiderablylowerthanseasonalratesofbetweenonceper7.
8e36.
0daysthatO'Connelletal.
(1988a)reportedfortheHadza.
Publisheddatafromotherchimpanzeeresearchsitesdonotallowcalculationofdailycarcass-encounterrates,butthesmallnumberofscavengingcasesimpliesthatratesareprobablysimilartothatatNgogo.
Thedensity,biomass,andspeciesdiversityofgrazingandbrowsingungulatesislowerinthehumidforeststhatconstitutemostchimpanzeehabitatsandinwoodlandssuchasGombethaninthedryer,moresea-sonalhabitatoftheHadzaand,presumably,intheapparentlywiderangeofhabitatsusedbyOldowanhominins(O'Connelletal.
,1988a;McGrew,1992;Stanford,1996;Schoeningeretal.
,2001;Plummer,2004).
Concomitantly,predatorguildsarelessdiverseinchimpanzeehabitats,andhumanshaveelim-inatedlargemammaliancarnivoresfromsome.
Additionally,decompositionisfasterinhumidforeststhanindryerwoodlandandsavannahabitats(Moore,1996;BoeschandBoesch-Acher-mann,2000;Roquetetal.
,2005).
Ediblecarcassesshouldthusbelesscommonandhaveshorterpersistenceincurrentlytypicalchimpanzeehabitatsthaninwoodlandsandsavannas.
Thesedifferencesincarcassabundancewereprobablygreaterinthepast,giventhatthehigherdiversityofbothungulatesandlargecarnivoresinPlio-PleistocenefaunasthaninrecentfaunasinAfricansavannahabitatspresumablyledtocorrespondinglyhigherdensitiesofpredatorsandmedium-to-large-sizedcar-casses(VanValkenburgh,2001).
Opportunitiesforconfronta-tionalscavengingmightthushavebeengenerallymorecommon,althoughtemporallyvariable,foranyhomininsabletoexploitthem(Blumenschine,1991;O'Connelletal.
,1988a).
Chimpanzeessometimesignorelarge,scavengeablemeatpackages,evenwhentheydonothavetoconfrontpredators;thisdistinguishesthemfromHadzaforagers,whotriedtoex-ploiteverypotentiallyprotablescavengingopportunitydur-ingO'Connelletal.
's(1988a)study.
WecannotknowhowoftenOldowanhomininspassedupopportunitiesforconfron-tationalscavenging,buttheymighthavedonesorelativelymoreoftenthanHadzahuntersdobecauseofgreatertechno-logicalconstraintsontheirabilitytodisplacelargecarnivoresfromkills(O'Connelletal.
,1988a;O'Connelletal.
,2002).
Chimpanzeeshavebeenobservedtoscavengeconfrontation-ally,butonlyagainstsmallerspeciesthatdonotpreyonthem(althoughadultmalebaboonsarepotentiallydangeroustoadultmalechimpanzees).
Observedcasesofscavengingkillsmadebyleopards,whichareknownpredatorsofchim-panzees(BoeschandBoesch-Achermann,2000),didnotin-volveconfrontations.
BoeschandBoesch-Achermann,(2000)notedthatchim-panzeesatTadonothuntduikerandseemnottorecognizethemaspotentialfood,despitetheirabundanceatthesiteandtheirinclusioninchimpanzeepreysetsatothersites.
Itispossiblethatmostchimpanzeesrecognizeanimalcarcassesasfoodonlyiftheyhavekilledtheanimals,seenotherchim-panzeekillthem,orseenotherchimpanzeeseatingthemdur-ingorafterhunts.
Chimpanzeesatallwell-documentedsiteshaveignoredatleastsomescavengingopportunitiesthatdonotfallintothesecategories(Table1);thisreluctancedistin-guishesthemfrommammaliancarnivoresinAfricansavannaecosystems(Plummer,2004).
InatleastfourcasesatGombeinwhichchimpanzeesscavengedpartiallyeatenredcolobuscarcasses,thescavengershadbeenatthehuntsandhadeithercaughttheprey,sawitcaught,fedfromit,orsawotherchim-panzeeseatingit(Goodall,1986).
RelevantinformationforothercasesandforMahaleandTaisnotavailable.
However,NgogoCases3and4andcasesofscavengingleopardkillsatMahale(Hasegawaetal.
,1983)conrmthatchimpanzeessometimesfeedonanimalsnotkilledintheirpresenceandnotkilledbyotherchimpanzees,includinganimalslargerthananytheycankill,althoughtheymayeatlittlefromthesecarcasses(e.
g.
,chimpanzeesatNgogocompletelyconsumedthecarcassinCase4,butatealmostnomeatinCase1)andindividualvariationinperceptionsofwhatisediblemayexist.
Alargeproportionofscavengingcaseshaveoccurredwhenchimpanzeesreturnedtositesofrecentmonkeyhuntstoeatmeatleftthere(Table1).
Eventhisisrare,probablybecausechimpanzeescompletelyconsumemostpreyimmediatelyaf-terhunts.
AtNgogo,individualsmaykeepmeatovernightaf-terhuntslateintheday,butrarelyleavelargeamountsofmeatatornearkillsites.
Case2wastheonlycaseofscavengingfromtheirownkills,andthemanypotentialconsumersatthemeat-eatingsessionwouldalmostcertainlyhaveeatenev-erythingedibleifnotforthedistractionbyneighbors.
OR'sdiscoveryofacarcassfarfromthehuntsitethefollowingmorningwasfortuitous.
GiventhatOldowanhomininsfacedcompetitionforfreshcarcasseswithmultiplelargeand130D.
P.
Watts/JournalofHumanEvolution54(2008)125e133dangerouscarnivorespecies(VanValkenburgh,2001;Plum-mer,2004),theirforagingtacticspresumablydidnotincluderetrievalofunconsumedmeatfrompreviouskillsites.
Chimpanzeesmayalsobelessinclinedtoexploitscaveng-ingopportunitiesthanarecarnivorousmammalsbecausetheylackcomparablephysiologicalbarrierstoinfectiousagentsincarrion(HamiltonandBusse,1978;cf.
PlummerandStanford,2000;Plummer,2004).
Inparticular,theymayfacehighrisksofcontractingzoonosesfromdeadordyingheterospecicsthattheyhavenotseenpredatorsattack.
Forexample,bothduikerandchimpanzeesaresusceptibletoEbola(Roquetetal.
,2005),andanthrax,botulism,andotherdiseasesaredi-rectlytransmissiblefromcarcasses(HamiltonandBusse,1978).
UseofleopardkillsatMahalesuggeststhatscavengingmightpreviouslyhavebeenmorecommoninhabitatsfromwhichleopardshavebeeneliminated(e.
g.
,Kibale).
However,Mahalechimpanzeesrespondedfearfullytothefreshcarcassofanadultbushpigkilledbyaleopardandthusignoredameatpackagefarlargerthantheirtypicalprey(Nishida,1994).
Additionally,researchershavenotobservedchimpan-zeesscavengingonleopardkillsatTa(whereleopardspreyonchimpanzees;Boesch,1991),althoughthismaybepartlybecauseTachimpanzeesdonothuntduiker,whicharethemajorpreyofleopards(BoeschandBoesch-Achermann,2000;ZuhberbuhlerandJenny,2002).
DosocialfactorscontributetotherarityofscavengingbychimpanzeesHuntingandespeciallymeateatinghavesocialimportance.
Allowingotherstohavemeatmaynotbecostlytohunterswhentheycapturepreytoolargeforasingleindivid-ualtoeat(e.
g.
,anadultmaleredcolobusmonkey)ortootherswhoobtainlargesharesofsuchkills,andcostsofharassmentbybeggarsmaysometimesoutweighthecostsofallowingthemtohavesomemeat(Gilby,2006).
Still,delayedreciproc-ityinmeatsharing,activesharing,andpreferentialsharingbe-tweenmaleallies(MitaniandWatts,2001,unpublisheddata)pointtowardsmeatsharingasameanstomaintainandstrengthensocialbondsbetweenmalesandtodevelopandmaintainal-liances.
Yetinsofarasmeatsharinghasthisfunction,itsef-fectsshouldnotdependonwhetherinitialmeatpossessorsobtainmeatviahuntingorscavenging.
Huntingbymalechimpanzeesmayinvolvecostlysignaling(HawkesandBliegeBird,2002).
Whilemeatsharingdoesnotincreasemalematingsuccess(MitaniandWatts,2001;Gilby,2006),huntingmonkeysisrisky,andmotivationtohuntandskillatpreycapturemaydisplaymalequality(HawkesandBliegeBird,2002)andsignalwillingnesstotakerisksandtoprovidetnessbenetstoothermales.
Suchsignalscouldinuencedecisionsaboutwhentoseekopportu-nitiestoengageinintergroupaggression(WattsandMitani,2001)andcouldinuencealliancestrategies(MitaniandWatts,2001).
Acquiringmeatthroughscavengingwouldnotprovidecomparablesignals.
Costly-signalingtheorymayindeedexplaincertainaspectsofhuntingandfoodsharingbyhumanhunter-gatherers,suchasHadzamen'sspecializationonhuntinglargepreythatbecomepublicgoods(HawkesandBliegeBird,2002)andturtlehuntingbyMeriammenwhoinvestconsiderabletimeandenergyinriskyhuntsbutdonatetheircatchtoothers(BliegeBirdetal.
,2001;Smithetal.
,2003).
However,theultimateexplanationforhuntingbychim-panzeesispresumablythatmeatisanextremelyvaluablenutritionalresource(MitaniandWatts,2001);thisremainstrueregardlessofitssource,solongaseatingitdoesnotimposetnesscosts(e.
g.
,becauseofdiseasetransmission).
Likewise,costlysignalingisrelevanttohuntingbyhumanspreciselybecausemeathassuchhighnutritionalvalue.
Hadzamenspecializeinhuntinglargegamedespitetheabsenceofdisproportionatenutritionalbenetstothemselvesandtheirfamilieswhentheysucceed,buttheyalsotypicallyexploitscavengingopportunities.
Confrontationalscavengingmayprovidesomescopeforcostlysignaling,butitsmainimportanceispresumablynutritional.
Itseemsunlikelythatchimpanzeeswouldfailtopursuesimilarnutritionalbenetssimplybecausepassivescavenginglackspotentialforcostlysignaling.
AtNgogo,chimpanzeescaptureonlyinfantandjuvenileredduikerandthelargestprey(adultmaleredcolobusandblack-and-whitecolobus)arefarsmallerthanthemodalsizeofungu-latespeciesusedbyBed1Olduvaihominins(Plummer,2004).
Scavengingcanincreasethesizerangeofcarcassesfromwhichchimpanzeesgainmeat(Table1):theadultredduikerinCase1waslargerthanallpreyitems(althoughRAandBIatealmostnoneofit),asweretheadultbushbuckscavengedatMahale(Hasegawaetal.
,1983).
However,itistooraretoproviderealnutritionalbenet.
Incontrast,scavengingcarcassesofmedium-sizedbovidsandlargermammalsprobablyaffordedanutritionallysignicantamountofmeattoOldowanhominins(PlummerandStanford,2000;Bunn,2001;Dominguez-RodrigoandPickering,2003;Plummer,2004).
Scavengingmighthavebeenmorecommoninmiombowoodlandhabitatsattheextremeofchimpanzeeecologicaltolerances(e.
g.
,Ugalla)thaninrainforestsandmorehumidwoodlands,butrecenthuman-inducedchangesinpredatorguildsmakeinvestigatingthispossibilitydifcult(McGrew,1992;Moore,1996).
LatePlioceneandearlyPleistocenehom-ininhabitatsexperiencedagradualtrendtowardsgreateropen-ness,withincreasedgrasslandafterabout2Ma,althoughvariationwasapparentlyextensive,withopengrasslands,bushlands,locallyforestedareas,andshrublandswithwet-landsallreconstructedforlocalitiesatwhichfossilsofHomoand/orParanthropushavebeenfound(ReedandRec-tor,2006).
Suchvariation,plusvariationinthelocaldensities,foragingstrategies,morphology,andgregariousnessoflargecarnivores,wouldhaveledtowidevariationinthefrequencyofscavengingopportunitiesforOldowanhominins(Plummer,2004),butopportunitiesshouldhavebeenmorecommonthanforchimpanzeesinforesthabitats.
TherecentdiscoveryofmiddlePleistocenechimpanzeefossils,datedtobetween5453kaand28412ka,intheKapthurinFormationoftheTugenHills,Kenya(McBreartyandJablonski,2005),castsdoubtonthenotionthatchimpanzeeevolutionaryhistoryhasbeendevoidofopportunitiestoscavengesimilartothoseavailabletoearlyHomoandofexposuretothesamepredatorguilds.
Thesefossilsshowthatthegeographicrangeof131D.
P.
Watts/JournalofHumanEvolution54(2008)125e133chimpanzeespreviouslyextendedintotheEastAfricanRiftValleywheretheywerepotentiallysympatricwithHomoinsemiarid,butlargelywooded,habitats(McBreartyandJablon-ski,2005).
Nevertheless,scavengingwouldprobablynothavebeenamajorcomponentofchimpanzeeforagingstrategiesinsuchacontextforseveralreasons.
Contemporaneoushomininsprobablycoulddrivelargecarnivoresfromkillsbythrowingrocksforcefullyandaccurately(VanValkenburgh,2001);chimpanzeeslackthisability,andtheydonotusetoolstohunt.
Additionally,wildchimpanzeeshaveneverbeenseenproducingsharp-edgedstoneakesofthesortthatwouldhaveallowedearlyhomininstoremovemeatfromcarcassesquickly,thenleaveacquisitionsitesandtherebyreducetheirownriskofpredation(Foley,1987;VanValkenburgh,2001);ancestralchimpanzeespresumablyalsolackedthisability.
Chimpanzeescanopenthelongbonesofcolobus-sizedpreywiththeirmolarstoextractmarrow(Teleki,1975;personalobservation),buthaveneverbeenseenusinghammerstonestoopenbones;withoutthisability,theycouldnothaveex-tractedmarrowfromlargercarcassestothesameextentasdidOldowanhominins.
Chimpanzeesinsemiarid,locallywoodedhabitatintheMt.
AssirikregionofSenegalareatriskofpredationbylionsandhyenasandarenotknowntoscavengefromeitherofthesecarnivores;thesamemightgenerallyhavebeentrueof''savanna''chimpanzeesincomparable,relativelyopenhabitats(Moore,1996;HuntandMcGrew,2002).
Stanford(2001:129)statedthat''meateatingpatternsbywildchimpanzeeslendlittlesupportforhypothesesforscav-engingasamajorcomponentoftheearlyhumandiet.
''Thisobservationmightberephrasedtostatethatdataonchimpan-zeecarnivoryhavelittlerelevancetohypothesesabouttheim-portanceofscavengingoncehomininsstartedtoincludemeatfrommedium-to-largemammalsintheirdiets,otherthantohighlightquestionsaboutdifferencesinhabitatpreferencesandotheraspectsofbehavioralecology,morphology,andcog-nition.
PassivescavengingmighthaveofferedlittlerewardtoOldowanhominins(Blumenschine,1991;Tappen,2001),anddeliberatesearchforconfrontationalscavengingopportunitiesmighthavebeenenergeticallyinefcientandrisky(Tappen,2001).
Nevertheless,iftheforagingstrategiesofsomePlio-cenehomininsincludedenoughscavengingtoyieldimportantnutritionalgains,andiftheycommonlyengagedinconfronta-tionalscavengingfromlargecarnivores,thenthesebehaviorswouldhavedistinguishedtheirbehavioralecologyfromthatofchimpanzees(PlummerandStanford,2000;Bunn,2001;Schoeningeretal.
,2001;Plummer,2004).
AcknowledgementsIthanktheMakerereUniversityInstitutefortheEnviron-mentandNaturalResources,TheUgandaWildlifeAuthority,andtheUgandaNationalCouncilforScienceandTechnologyforpermissiontoworkatNgogo.
IamindebtedtoJohnKase-neneandGilIsibirye-BasutaforpermissiontoworkattheMakerereUniversityBiologicalFieldStationandforalltheirsupportovertheyears.
JohnMitaniandJerryLwangahavebeenoutstandingcolleagues,collaborators,andfriendsatNgogoandhavecontributedgreatlytomyunderstandingofhuntingandmeatsharingbychimpanzeesthere.
FieldworkatNgogoispossibleonlywiththeexpertassistanceofGod-freyMbabazi,AdolphMagoba,LawrenceNdagezi,andAlfredTumusiime.
MyresearchatNgogohasbeensupportedbytheL.
S.
B.
LeakeyFoundation,theNationalGeographicSociety,PrimateConservation,Inc.
,andYaleUniversity.
BillKimbel,ManuelDominguez-Rodrigo,andananonymousreviewerpro-videdextremelyhelpfulcommentsonanearlierversionofthismanuscript.
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133D.
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Watts/JournalofHumanEvolution54(2008)125e133
WattsDept.
ofAnthropology,YaleUniversity,P.
O.
Box208277,NewHaven,CT06511-8277,USAReceived28February2007;accepted3July2007AbstractChimpanzeesregularlyhuntavarietyofpreyspecies.
However,theyrarelyscavenge,whichdistinguisheschimpanzeecarnivoryfromthatofsomemodernhunter-gatherersand,presumably,atleastsomePlio-Pleistocenehominins.
Iuseobservationsmadeoveran11-yearperiodtodocumentallknownopportunitiesforscavengingencounteredbychimpanzeesatNgogo,KibaleNationalPark,Uganda,anddescribeallcasesofscavenging.
Ialsoreviewdataonscavengingfromotherchimpanzeeresearchsites.
ChimpanzeesatNgogoencounteredscavengingoppor-tunitiesonlyaboutonceper100daysandatemeatfromscavengedcarcassesonlyfourtimes.
Scavengingopportunitiesarealsorareatothersites,evenwhereleopardsarepresent(Mahale,Ta,Gombe),andscavengingofleopardkillsisknownonlyfromMahale.
Feedingonpreythatchimpanzeeshadhuntedbutthenabandonedisthemostcommonformofscavengingreportedacrossstudysites.
Forexample,severalindivid-ualsatNgogoatemeatfromapartiallyconsumedredcolobuscarcassabandonedafterahuntthepreviousday.
SuchbehaviorprobablywasnotcommonamongOldowanhominins.
Ngogodataandthosefromothersitesalsoshowthatchimpanzeessometimeseatmeatfromcarcassesofpreythattheydidnotseekilledandthatwerenotkilledbychimpanzees,andthatscavengingallowsaccesstocarcasseslargerthanthoseofanypreyitems.
However,chimpanzeesignorerelativelymanyopportunitiestoobtainmeatfromsuchcarcasses.
Scavengingmayberarebecausefreshcarcassesarerare,becausetheriskofbacterialinfectionsandzoonosesishigh,andbecausechimpanzeesmaynotrecognizecertainspe-ciesaspotentialpreyorcertainsizeclassesofpreyspeciesasfoodsources.
Itsminimalnutritionalimportance,alongwiththeabsenceoftech-nologytofacilitateconfrontationalscavengingandrapidcarcassprocessing,apparentlydistinguisheschimpanzeeforagingstrategiesfromthoseofatleastsomeOldowanhominins.
2007ElsevierLtd.
Allrightsreserved.
Keywords:Chimpanzees;Meateating;Oldowanhominins;ScavengingIntroductionWildchimpanzeesregularlyhuntandconsumeavarietyofvertebrateprey,andtheyeatmoremeatthandomostorallothernonhumanprimates.
Detaileddataonhuntingandmeatconsumptionhavebeencollectedatmultiplesites,espe-ciallyatGombe(Goodall,1986;Stanford,1998),Ta(BoeschandBoesch,1989;BoeschandBoesch-Achermann,2000),Mahale(Hosakaetal.
,2001),andNgogo(MitaniandWatts,1999,2001;WattsandMitani,2002).
Chimpanzeessometimesopportunisticallypursuesolitaryprey,butmoreoftenconductgrouphuntsofgregariouspreyinwhichtheymaydeploycom-plex,cooperativeprey-capturetactics(BoeschandBoesch,1989;BoeschandBoesch-Achermann,2000).
Theyalsocom-monlysharemeataftersuccessfulhunts.
Comparativedataonchimpanzeehuntingandmeatconsumptionarevaluableforinvestigationsoftheimportanceofhuntingandmeateatinginhumanevolution(e.
g.
,Moore,1996;Stanford,1996,2001;Kaplanetal.
,2001)andtheevolutionaryecologyoffoodsharingbyhumans(e.
g.
,Winterhalder,2001).
Incontrast,reportsofscavengingbychimpanzeesarerare.
Inthispaper,IsummarizepublisheddataonscavengingbychimpanzeesandpresentnewdatafromNgogo,KibaleNa-tionalPark,Uganda.
IdiscussthesedatawithreferencetothedebateovertherelativeimportanceofhuntingandE-mailaddress:david.
watts@yale.
edu0047-2484/$-seefrontmatter2007ElsevierLtd.
Allrightsreserved.
doi:10.
1016/j.
jhevol.
2007.
07.
008Availableonlineatwww.
sciencedirect.
comJournalofHumanEvolution54(2008)125e133scavenginginthesubsistencestrategiesofPlio-Pleistocenehominins.
ChimpanzeesatNgogohuntfrequentlyandareun-usuallysuccessfulpredatorsofredcolobusmonkeys(MitaniandWatts,1999,2001;WattsandMitani,2002),buttheyig-noremostoftheirrareopportunitiestoscavenge,althoughtheyhavebeenobservedtoeatpartsoffourcarcasses.
Thisseemstypicalofchimpanzeesinforesthabitats,andthecon-tributionofscavengingtochimpanzeesubsistenceisminimal.
ScavengingbymodernhumansandPlio-PleistocenehomininsThesubsistencestrategiesofsomemodernhumanhunter-gatherersincludescavenging.
Thebest-studiedexampleistheHadza,whomonitorlargecarnivoresandvulturestolearnofscavengingopportunitiesthattheyexploiteitherpassivelyorbyconfrontingthecarnivores,includinglions,andstealingtheirkills(O'Connelletal.
,1988a;Bunn,2001).
Ofallthemedium-to-large-sizedanimalcarcassesthatagroupofHadzaforagersacquiredduring14monthsin1985e1986,20%werescavenged(O'Connelletal.
,1988a).
Confrontationalscaveng-ingledtoacquisitionofcarcassesthatwereover75%completeinveof11scavengingcasesandoccasionallypro-videdlargeamountsofmeat.
However,seasonalvariationincarcass-encounterratesandvariationinthesizeandcomplete-nessofcarcassesledtohighvarianceinreturnsfromscaveng-ingandmadescavengedanimaltissueonlya''windfallresource.
''Inaddition,passivescavengingyieldedlittlemeat(O'Connelletal.
,1988a).
Extensivedebatesurroundsthequestionsof(1)howimpor-tantscavengingwas,comparedtohunting,inPlio-Pleistocenehomininsubsistencestrategiesand(2)wheninthecourseofhumanevolutionmeatbecameacriticaldietarycomponent(re-viewedbyDominguez-Rodrigo,2002;O'Connelletal.
,2002;Dominguez-RodrigoandPickering,2003;Plummer,2004).
Thisdebatelargelyinvolvescontrastinginterpretationsofre-sultsfromanalysesoffaunalassemblagesfromOldowansitesthatshowhominin-inducedmodication.
Italsoinvolvesdis-agreementabouttherelativecontributionsoffoodsofanimalorplantorigintothedietsofmodernhumanhunter-gatherersand,concomitantly,(1)therelativeimportanceofmen'sandwomen'scontributionstohunter-gatherersubsistence;(2)theroleofhuntingandmeatsharinginmalereproductivestrategies;and(3)therelevanceofethnographicexamplestore-constructionsofearlyhomininbehavioralecology.
Iwillonlybrieyreviewsomeaspectsofthisdebatetowhichchimpanzeedataarepotentiallyrelevant.
IfanyOldowanhomininsfre-quentlyscavenged,andscavengingcontributedsubstantiallytotheirdiet,thenthiswouldhavebeenamajordifferenceinbe-havioralecologyfromthatofextantchimpanzees.
Inaddition,ifscavenginghadonlyminornutritionalimportance,butcon-frontationalscavengingofcarnivorekillswasimportantasamalematingtactic(O'Connelletal.
,2002),thenthiswouldraisetheissueofwhether,andifso,how,meatsharingentersintothereproductivestrategiesofmalechimpanzees.
DeterminingthetimingofaccesstocarcassesandhowmuchmeattheycouldhaveprovidediscrucialtoassessinghowOldowanhomininsassembledmodiedanimalbones.
Manykindsofdataarerelevanttosuchdeterminations,in-cludingthoseonthefrequencyandlocationofstone-toolcutmarksandcarnivoretoothmarks,onbone-fracturepatterns,onskeletal-partproles,andonhowextantcarnivoresandcontemporaryhumanforagers(e.
g.
,theHadza:O'Connelletal.
,1988b;Bunn,2001)modifycarcasses.
Extensiveanaly-sesofungulateremainsfromseveralBedIsites(especiallyFLKIZinjanthropus)atOlduvaiGorge,Tanzania,mostofwhicharefrompreyofsizeclasses2or3(adultbodyweight40e300kg)andwhichdatetoabout1.
75millionyearsago(Ma),haveledtocontrastinginterpretations:(1)homininshadonlylateaccesstocarcassesandprobablyonlyextractedmarrowfrombonesmostlydeeshedrstbycarnivores(Blu-menschine,1991),or(2)theyregularlyhadearlyaccesstocar-casses,consistentwitheitherhuntingortheftofcarcassesfromlargecarnivores(BunnandKroll,1998;Dominguez-Rodrigo,1997,1999;Bunn,2001;O'Connelletal.
,2002;Bunn,2006).
DatafromseveralotherPliocenesitesareconsistentwithearlyaccesstocarcasses.
Inparticular,highfrequenciesofcutmarksonlong-bonemidshaftsandothermeatyskeletalelements,combinedwithlowfrequenciesofcar-nivoretoothmarksontheseelements,pointtoearlyaccessatOlduvaisiteFxJj50(Dominguez-Rodrigo,2002),Peninj(Dominguez-Rodrigo,2002),andperhapsatGona(Domi-nguez-Rodrigoetal.
,2005).
InarecentreanalysisofdatafromFLK-Zinjanthropusandcritiqueofearlieranalyses,Dom-inguez-RodrigoandBarba(2006)concludedthathomininsalsohadearlyaccesstothecarcassesofmedium-sizedbovidsaccu-mulatedatthissite.
Earlycarcassaccessdidnotnecessarilyentailhunting.
O'Connelletal.
(2002)agreedthatbone-modicationdataandactualisticpatternsshowearlyaccesstomanycarcasses,butcontendedthatlackofprojectiletechnologypreventedPlio-Pleistocenehomininsfromhuntinglargegame,andthattheygainedearlyaccessbyaggressivelyconfrontingpredatorsatrelativelyfreshkills.
Likewise,Stiner(2002:6)arguedthat,whilesomehuntingofsmallerpreyprobablyoccurred,largemammals''wouldhavebeendifculttokillwithOldowanorearlyAcheuleantechnology,''althoughshealsonotedthattheinitialoccupationofpartsofEurasiabyhomininsthatpossessedsuchtechnologydependedonforagingstrategiesthatincludedcarnivory.
However,Dominguez-Rodrigoetal.
's(2001)plant-residueanalysisofAcheuleantoolsfromPeninjprovidedevi-denceforwoodworkingat1.
5Ma,fromwhichtheyconcludedthatHomoergasterhadtheabilitytoproducespearsthatcouldhavebeenusedforhunting.
O'Connelletal.
(2002)alsoarguedthathighcarnivoretooth-markcountsonbonesfromOldowansitesindicateaggressivescavengingofcarcassesalreadyde-eshedtovaryingdegrees,nothunting.
Dominguez-Rodrigo(2002,2003;cf.
Dominguez-RodrigoandPickering,2003)counteredbynotingthatthehighfrequencyofcutmarksonboneswherecarnivoresstartdeeshing(e.
g.
,upper-limblongbones,pelves),andespeciallythehighfrequencyofcutmarksonupper-limblong-boneshafts,resemblesthepatternthatre-sultswhenhumansinitiatethedeeshing,ashappenswhentheyhavehuntedtheanimals.
126D.
P.
Watts/JournalofHumanEvolution54(2008)125e133SeveralresearchershavearguedthatmeatobtainedfromconfrontationalscavengingwasnotamajorcomponentofOldowanhominindiets,foravarietyofreasons.
Forexample,Blumenschine(1991)arguedthattherisksoftryingtostealfreshcarcassesfromlargecarnivoreswouldhavebeenex-tremelyhigh(withtheexceptionofcachedleopardkills)andthattheoccurrenceofscavengingopportunitieswouldhavevariedgreatlyonaseasonalbasis.
Othershavearguedthatconfrontationalscavengingwouldhaveoccurredonlyifthehomininshadconsiderablenumericalsuperiorityovertheircarnivorecompetitorsand/oriftheycouldreducetheriskssuf-cientlybyusingweapons(Shipman,1986;O'Connelletal.
,1988a,b;VanValkenburgh,2001).
BasedontheirstudyofHadzascavenging,Hawkesetal.
(1988a)assertedthathighvarianceinthetimingofopportunitiesandinpossiblereturnswouldhavemadeconfrontationalscavengingatbestasourceofoccasionalwindfallsforOldowanhominins,notaregularactivity.
AstheirexampleofHadzascavengingofanearlyin-tactgiraffecarcassindicated,however,returnsmightsome-timeshavebeenextremelyhigh.
AgainrelyingonHadzadata,O'Connelletal.
(2002)arguedthatconfrontationalscav-engingbymalesservedmainlyascostlysignalingthatgavefemalesaccurateinformationaboutmatequality,ratherthanasanimportantsourceofnutrients.
Intheirview,meatwasanunreliablefoodsourceanditsdietaryimportancesecondarytothatofplantfoods,particularlyundergroundstorageorgans.
However,othershavecautionedagainsttakingtheHadzaeitherasgeneralrepresentativesofmodernhunter-gatherersorasanalogiesforOldowanhominins(e.
g.
,Kaplanetal.
,2001;Dominguez-Rodrigo,2002,personalcommunication).
Areviewoftheethnographicrecordsupportstheclaimthatmostmodernhunter-gatherersgainedmostoftheircaloriesfrommeat,andthusfromhunting,amostlymaleactivity(Kaplanetal.
,2001).
Dominguez-Rodrigo(2002;c.
f.
Dominguez-RodrigoandPickering,2003)attributedmajorim-portancetomeatinOldowanhomininsubsistence,butarguedthatmostofitcamefromhunting,notconfrontationalscavenging.
Plummer's(2004)preliminaryanalysisofprey-sizedistributionandevidenceforbutcheryandcarnivoreac-tivityatthe2-million-year-oldsiteofKanjerahelpstoshiftthedebateawayfromalongstandingfocusonlargegame.
Amongotherlinesofevidence,highfrequenciesofstone-toolcutmarks,lowfrequenciesofcarnivoretoothmarksonlimb-boneshafts,andhighrepresentationofjuveniles(usuallycompletelydestroyedbycarnivores)indicatethathomininsregularlyhuntedsmall(10a17!
118Scavengingcases10b7c7b4Casesfromearlierredcolobushunts5e9d30e1dScavengenonpreyspeciesNoNoYes(bushbuck)NoExtendpreysizerangeNoNoYesYesIgnorelargepreyYes(bushpig,bushbuck)(duiker)eYes(bushpig)YesStealkillsbyotherpredatorsBaboonCrownedeagleLeopardCrownedeagleConfrontationalcarcasstheftYesYesNoNoFeedoncarcasseswhenpredationunseenNoNoYesYesSources:Gombe:Goodall(1986),Mulleretal.
(1995),W.
Wallauer(personalcommunication);Ta:BoeschandBoesch-Achermann(2000);Mahale:Hasegawaetal.
(1983),Nishida(1994);Ngogo:thisstudy.
aNotincludingcarcasstheftsfrombaboonsorencounterswithbaboonseatingmeat(Goodall,1986;numberofcasesnotreported).
bChimpanzeesobtainedonecarcassfromhumansatGombe(Goodall,1986)andtwofromhumansatMahale(Hasegawaetal.
,1983).
cDoesnotincludethreecasesinwhichchimpanzeesstolelivingpreyfromcrownedeagles.
dFivedeniteandfourprobableGombecases;informationnotprovidedforonecaseatMahale.
eChimpanzeesatTaignoreduikerkilledbyleopards,butalsodonothuntduiker.
127D.
P.
Watts/JournalofHumanEvolution54(2008)125e133casesofscavenginghavebeenobservedatGombesinceGoodall's(1986)review(W.
Wallauer,personalcommunica-tion).
Hasegawaetal.
(1983)reportedfourcasesofscaveng-ingatMahale,twoofadultbushback(Tragelephusscriptus)probablykilledbyleopards(onenearlyintact,thesecondfreshbutmostlyeaten)andtwoofblueduiker(Cephalophuscalli-pyga)carcassesthatresearchershadhandled.
Nishida(1994)laterreportedthreemorecases:oneblueduiker,onered-tailedmonkey(Cercopithecusascanius),andoneredcolobus(Pilio-colobustephrosceles).
Basedonobservationsovernearlytwodecades,BoeschandBoesch-Achermann(2000)reportedonlysevencasesatTa,includingthreeoffeedingonremainsofmonkeyskilledbychimpanzeesoneortwodayspreviouslyandfouroffeedingonmonkeyskilledbycrownedeagles(Ste-phanoaetuscoronatus).
Chimpanzeesalsostolewoundedredcolobusfromeaglesthreetimesandencountered,butignored,tenfreshcarcassesofseveralmammalianspecies,althoughonlyoneofthese(sootymangabeys,Cercocebusatys)wasaknownpreyspecies.
Reynolds(2005)reportedonecasefromBudongo,inwhichchimpanzeesfedonthecarcassofaninfantbluemonkey(Cercopithecusmitis)thathadbeenkilledandpartiallyconsumedbyanadultmalebluemonkey;chimpanzeesoccasionallypreyonbluemonkeysatthissite.
ScavengingbychimpanzeesatNgogoMethods:studysiteandsubjectsNgogoisinthecenterofKibaleNationalPark,inwesternUganda.
Thestudyarea,atanaltitudeofabout1400m,receives1700e1800mmofrainannually,mostlyfromMarchthroughMayandmid-AugustthroughDecember.
Itcoversabout30km2ofmixed,mature,andregeneratingforestthatistransi-tionalbetweenlowlandandmontaneevergreenforestandincludesother,minor,vegetationtypes(Butynski,1990;Struhsaker,1997;Lwangaetal.
,2003).
Threesmall-to-medium-sizedungulatesarecommon(Struhsaker,1997):redduiker(Cephalophusmonticola),blueduiker,andbushpig(Potomochoerusporcus).
Bushbuckarealsopresent,butarelesscommon.
Sevenotherdiurnalprimatespeciesarepresent,withred-tailedmonkeysandgrey-cheekedmangabeys(Lophocebusalbigena)beingthemostcommon(Mitanietal.
,2000).
Thechimpanzeecommunityhasbeencontinuouslyob-servedsincemid-1995andisthelargestknown;its140e150membershaveincluded22e28adultmalesand14e16adolescentmales(Watts,1998;MitaniandWatts,2005).
Thechimpanzeespreyonatleast11vertebratespecies,in-cludingsixdiurnalprimatesandthethreecommonungulates(althoughtheydonothuntadultbushpigsoradultreddui-ker).
Thepredationdatabaseincludesover650killsduringmorethan270hunts(WattsandMitani,2002,unpublisheddata).
Redcolobusmonkeysarebyfarthemostcommonprey;about80%ofredcolobushuntsresultinkills,andchimpanzeesmakemultiplekillsofredcolobusinmosthunts.
IobservedchimpanzeesatNgogoforover10,000hoursbe-tween1995and2006.
Irecordedallhuntsandpreycaptures,collecteddataonmeatsharing,andnotedallencounterswithfreshprey-speciescarcassesnotkilledbychimpanzeesandallcasesofscavenging.
ResultsChimpanzeesencounteredonlyeightrecentlydead,nonde-composedcarcassesofpreyspeciesandfoundoneindividualofapreyspeciesdyingfromunknowncauses.
Thus,theen-counterratewithscavengeablecarcasseswasonlyonceperca.
98days,assuminga13-houractivityperiod.
Thisincludestwocasesinwhichadultmalebaboonswereeatingredduikerthattheyhadjustkilled;thechimpanzeesshowednointerestintryingtostealthesecarcasses.
Intwoothercases,chimpan-zeesencountereddeadadultredduikerthattheydidnotcon-sume.
Intherstofthesecases,threemalechimpanzeesignoredthecarcass,andinthesecond,anadultfemalewithaninfantandanadolescentfemalebrieyinspectedthecar-cass,butdidnottouchit.
Thefourscavengingepisodes,andoneinteractionbetweenanadultmalechimpanzeeandadyingblack-and-whitecolobusmonkey(Colobusguereza),aredetailedbelow.
Case1:adultfemaleredduikerAt0920honSeptember26,2002,adolescentmalesRAandWAandadolescentfemaleBIfoundadeadadultfemaleredduiker.
Itsabdomenhadanarrow6-cmopeningandsomeviscerahadbeenremoved;nootherwoundswerevisibleandthecarcasswasotherwiseintact.
Thelimbswereslightlyex-ible,butmanyieswereswarmingaroundthecarcass.
WAandBIinspecteditcloselyandsniffedit,butdidnottouchit.
RAapproachedit,sniffedandvisuallyinspecteditandpokedatitsventrum.
Hepulledandtwistedoneoftheduiker'slegswithbotharmsforfourminutesasiftryingtodisarticu-lateit,pokedattheopeningintheabdomenagainandsniffedhisnger,thendraggedthecarcass3mandpulledandtwistedthelegforanotherveminutes.
Finallyhebitintothelegsev-eraltimes,piercingtheskinjustabovethekneeandremovingasmallamountofmeat,whichheate;hedidnotbiteintothemeatierhipregion.
Hesatnexttothecarcassforseveralmoreminutesandthenleftthesite.
WAfollowedhim,butBIthenbrieypulledandtwistedoneleg,bitintoit,andremovedasmallamountofmeat,whichsheate.
Shethenfollowedthemales.
SofarasIwasaware,nootherchimpanzeesencounteredthecarcass.
Case2:adultfemaleredcolobusAt1645honDecember10,2002,15adultmalechimpan-zees,veadolescentmales,12adultfemales(sevenwithin-fants),twoadolescentfemales,andninejuvenilesencounteredaredcolobusgroup.
Theystartedtohuntthemonkeysat1710handmadevekills(oneadultfemale,twosubadults,andtwoinfants)betweenthenand1755h.
Onemoreadult128D.
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Watts/JournalofHumanEvolution54(2008)125e133malechimpanzeeandoneadolescentmalearrivedduringthesubsequentmeat-eatingsession.
At1820h,chimpanzeescreamswereaudiblefromthenorth;thecallersbelongedtoaneighboringcommunity.
TheNgogomalesbecamehighlyexcitedandrushedinthatdirec-tion.
Atleasttwomaleswerecarryingmeat(thetwolargestcarcasses),butdroppeditastheyapproachedthecallers.
At1842h,themalesstoppedandlistenedonthesouthernedgeofasmallvalley;chimpanzeescalledfromthenorthsideofthevalleyat1845h.
TheNgogomalesexcitedlybutsilentlyapproached,andat1858htheychargedtheotherchimpan-zees,whoscreamedandednorth.
Malescontinuedtodisplayuntil1915h,thenmovedsouthandnestedintheareaofthehunt.
Onthenextmorning,11oftheadultmales,threeadoles-centmales,oneorphanedjuvenilemale,andseveralfemales,somewithdependentoffspring,returnedtothesouthernedgeofthevalleywherethemaleshadattackedtheneighborsthepreviousevening.
At1303h,allthemalesplusoneoldfemalestartedapatroltothenorthandeast,duringwhichtheytrav-eledseveralkilometerswithoutencounteringneighbors.
At1205h,beforethepatrolstarted,adultmaleORfoundmostofanadultredcolobuscarcass,presumablythefemalekilledthepreviousafternoonbyHA,ontheground.
Hepickeditup,carriedit20m,thenclimbedatreeandstartedtoeatmeat.
Meanwhile,adultmaleAYfoundpartofalimb,appar-entlyfromthesamemonkey,andstartedtoeatmeat.
ORsharedmeatwithtwoadolescentmales.
One(DX)brieyre-trievedthecarcassandatemoremeatwhenORdroppeditat1240tofollowothermalesacrossthevalley.
Thesefourmalestreatedthecarcasslikeafreshkill:nonehesitatedtoeatmeat,althoughtheydidnoteatallofit.
Nootherchimpanzeesshowedinterestinobtainingmeat.
Case3:subadultred-tailedmonkeyOnthemorningofJuly12,2005,14adultmales,6adoles-centmales,anestrousfemaleandheradolescentandjuveniledaughters,andatleastsevenotheradultfemaleswithveju-venilesandfourinfantswereeatingChrysophyllumwarwickiifruitinseveraltrees.
At1310h,youngadultmaleGTZfoundadeadsubadultmalered-tailedmonkeyonthegroundbeneathoneofthesetrees.
Itappearedfreshditslimbswereexibledandhadnoobviouswounds(althoughthoseinictedbycrownedeaglesmaybevisibleonlyoncloseinspection;T.
Struhsaker,personalcommunication).
GTZgraspeditstailanddraggedit10masadolescentmaleMUwatched,thenstoppedandbitintooneofitshipjoints.
Afterseveralminutesofbitingandpullingatthejoint,hetoreoffonelegandthetail,carriedtheseupatree,andstartedtoeatmeat.
Noonejoinedhimorbeggedformeat.
AdultmaleLO,whoarrivedwhileGTZwasdisarticulatingthelegandtail,draggedtherestofthecarcass20malongtheground,thensatandlookedatitfor3min,butatenomeat.
Nootherchimpanzeesinves-tigatedthecarcass.
GTZwasstilleatingmeatat1400h,whenfouradultmalesandseveralfemalesandtheirdependentoffspringmovedsouthandencounteredaredcolobusgroupabout1kmfromthescavengingsite.
Onemaleclimbedtowardtheredcolobus,butthemonkeysed.
GTZ,stillcarryingpartofthered-tailedmonkeycarcass,arrivedwithseveralothermalesat1510h.
Heclimbedabout12mupaPseudospondiasmicrocarpatreeandresumedeatingmeat.
Anadolescentfemalejoinedhimandbeggedunsuccessfullyformeat.
At1605,thechimpanzeeshuntedblack-and-whitecolobusinthesameareaandadultmaleMOkilledajuvenile,halfofwhichanotheradultmalestole.
GTZdidnotparticipateinthehuntorjoinbegging/meat-sharingclustersthatformedaroundthetwomeatpossessors.
Hedroppedtheremainsofthered-tailedmonkeysometimebetweenthenand1700h.
Case4:subadultredduikerAt1215honApril4,2006,alargepartyofchimpanzeesencounteredthecarcassofasubadultredduikerlyingonthegroundinaswampyvalley.
Theduikerappearedtobefreshlydeadandmighthavebeenkilledbyacrownedeagle.
Sixadultmalesplusseveralyoungadolescentmalesandmanyfemales,juveniles,andinfantscrowdedaroundthecar-cassandlookedatit.
Noneoftheadultstoucheditatrst,butseveraljuvenilestouched,sniffed,andhitit.
JuvenilemaleBNstartedtodragitaway;thispromptedalphamaleBTtochargethroughthemiddleoftheassembledgroup.
Mostofthechim-panzeesed,andmanyothersalsochargedatorotherwiseen-gagedinaggressiveinteractions.
Duringthismelee,BNletgoofthecarcassandayoungadolescentfemaledraggeditintothickvegetation,butthenalsoletgoofit.
Atsomepoint,adultmalePAseizedthecarcassandcarrieditupaPseudospondiasmacrocarpatree,wherehestartedtoeatit.
Sixotheradultmales,oneadolescentmale,andonejuvenilemalefollowedandstartedtobegformeat.
PAwasreluctanttoshareandman-agedtopreventallbutalphamaleBTfromobtainingmeatwhilehefedontheduikerforoveranhour.
BTleftduringthistime,buttheothermalesbecameincreasinglyagitated,andnally,adultLOchargedatPAandprecipitatedanothermelee,duringwhichhestolethecarcassfromPA.
AdultmaleMORthenstoleitfromhim.
MORsharedmeatwiththreeotheradultmalesandanadolescentfemale,andrefusedbeggingattemptsbyafourthadultmale.
Oneotheradultmaleandayoungadolescentmaleobtainedscrapsofmeatfromtheground.
Theentiremeat-eatingsessionlastedalmostfourhoursandthechimpanzeesconsumedtheentirecarcass.
Case5:subadultmaleblack-and-whitecolobusAt1600honJuly1,2002,alphamaleBTlefteightotheradultmales,twoadolescentmales,andonefemalewithajuve-nileandinfanttoforagealone.
At1610h,heheardalarmcallsfromagroupofblack-and-whitecolobusmonkeys;helookedtowardsthemonkeys,thenrushedoverandsatbelowthem.
Suddenlyhespottedalargesubadultmalemonkeyontheground,alivebutnotmoving.
Itwasnotobviouslywounded,andmighthavefallenbecauseofanaccident,becauseofag-gression,becauseitwasill,orbecauseofacrownedeagle129D.
P.
Watts/JournalofHumanEvolution54(2008)125e133attack.
BTdraggedthemonkey30mbyitstail,thrashingitagainstthegroundseveraltimes;itresistedineffectivelybygrabbingatplantstems.
Afterstoppingtositandlookatit(withoutreleasinghisgrip),hedraggeditanother10m,thrasheditagainstthegroundagain,andcarrieditupaPseu-dospondiasmicrocarpatree,whereheplaceditonalargeboughandsatnexttoit.
Hehitthemonkeyagainsttheboughseveraltimes,sniffeditandinspecteditvisually,andputhismouthonitshead,butdidnotbiteit.
Hethencarriedittoasecondbough,putitdown,andsatlookingcloselyatit.
Themonkeywasnolongermovingandappeareddead.
Tenminutesafterherstclimbedthetree,BTdescendedandmovedoffwithouthavingeatenanymeat;heleftthemonkeyinthetree.
DiscussionTheseobservationsaddtothesmallbodyofdatashowingthatwildchimpanzeesinlowlandevergreenmoistforests(Ta),mid-altitudeevergreenmoistforest(Kibale,Budongo),anddryer,moreseasonalforests(Mahale)andwoodland(Gombe)scavengevariousvertebratespeciesonwhichtheyalsoprey.
However,scavengingisrare,especiallycomparedtoratesofpredationonsomeofthesamespecies,andchim-panzeesdeclinealargeproportion,andmaybethemajority,ofscavengingopportunities(Table1).
Oneobviousreasonfortherarityofscavengingistherarityofopportunities.
Theestimateofoneencounterper98daysatNgogoisconsiderablylowerthanseasonalratesofbetweenonceper7.
8e36.
0daysthatO'Connelletal.
(1988a)reportedfortheHadza.
Publisheddatafromotherchimpanzeeresearchsitesdonotallowcalculationofdailycarcass-encounterrates,butthesmallnumberofscavengingcasesimpliesthatratesareprobablysimilartothatatNgogo.
Thedensity,biomass,andspeciesdiversityofgrazingandbrowsingungulatesislowerinthehumidforeststhatconstitutemostchimpanzeehabitatsandinwoodlandssuchasGombethaninthedryer,moresea-sonalhabitatoftheHadzaand,presumably,intheapparentlywiderangeofhabitatsusedbyOldowanhominins(O'Connelletal.
,1988a;McGrew,1992;Stanford,1996;Schoeningeretal.
,2001;Plummer,2004).
Concomitantly,predatorguildsarelessdiverseinchimpanzeehabitats,andhumanshaveelim-inatedlargemammaliancarnivoresfromsome.
Additionally,decompositionisfasterinhumidforeststhanindryerwoodlandandsavannahabitats(Moore,1996;BoeschandBoesch-Acher-mann,2000;Roquetetal.
,2005).
Ediblecarcassesshouldthusbelesscommonandhaveshorterpersistenceincurrentlytypicalchimpanzeehabitatsthaninwoodlandsandsavannas.
Thesedifferencesincarcassabundancewereprobablygreaterinthepast,giventhatthehigherdiversityofbothungulatesandlargecarnivoresinPlio-PleistocenefaunasthaninrecentfaunasinAfricansavannahabitatspresumablyledtocorrespondinglyhigherdensitiesofpredatorsandmedium-to-large-sizedcar-casses(VanValkenburgh,2001).
Opportunitiesforconfronta-tionalscavengingmightthushavebeengenerallymorecommon,althoughtemporallyvariable,foranyhomininsabletoexploitthem(Blumenschine,1991;O'Connelletal.
,1988a).
Chimpanzeessometimesignorelarge,scavengeablemeatpackages,evenwhentheydonothavetoconfrontpredators;thisdistinguishesthemfromHadzaforagers,whotriedtoex-ploiteverypotentiallyprotablescavengingopportunitydur-ingO'Connelletal.
's(1988a)study.
WecannotknowhowoftenOldowanhomininspassedupopportunitiesforconfron-tationalscavenging,buttheymighthavedonesorelativelymoreoftenthanHadzahuntersdobecauseofgreatertechno-logicalconstraintsontheirabilitytodisplacelargecarnivoresfromkills(O'Connelletal.
,1988a;O'Connelletal.
,2002).
Chimpanzeeshavebeenobservedtoscavengeconfrontation-ally,butonlyagainstsmallerspeciesthatdonotpreyonthem(althoughadultmalebaboonsarepotentiallydangeroustoadultmalechimpanzees).
Observedcasesofscavengingkillsmadebyleopards,whichareknownpredatorsofchim-panzees(BoeschandBoesch-Achermann,2000),didnotin-volveconfrontations.
BoeschandBoesch-Achermann,(2000)notedthatchim-panzeesatTadonothuntduikerandseemnottorecognizethemaspotentialfood,despitetheirabundanceatthesiteandtheirinclusioninchimpanzeepreysetsatothersites.
Itispossiblethatmostchimpanzeesrecognizeanimalcarcassesasfoodonlyiftheyhavekilledtheanimals,seenotherchim-panzeekillthem,orseenotherchimpanzeeseatingthemdur-ingorafterhunts.
Chimpanzeesatallwell-documentedsiteshaveignoredatleastsomescavengingopportunitiesthatdonotfallintothesecategories(Table1);thisreluctancedistin-guishesthemfrommammaliancarnivoresinAfricansavannaecosystems(Plummer,2004).
InatleastfourcasesatGombeinwhichchimpanzeesscavengedpartiallyeatenredcolobuscarcasses,thescavengershadbeenatthehuntsandhadeithercaughttheprey,sawitcaught,fedfromit,orsawotherchim-panzeeseatingit(Goodall,1986).
RelevantinformationforothercasesandforMahaleandTaisnotavailable.
However,NgogoCases3and4andcasesofscavengingleopardkillsatMahale(Hasegawaetal.
,1983)conrmthatchimpanzeessometimesfeedonanimalsnotkilledintheirpresenceandnotkilledbyotherchimpanzees,includinganimalslargerthananytheycankill,althoughtheymayeatlittlefromthesecarcasses(e.
g.
,chimpanzeesatNgogocompletelyconsumedthecarcassinCase4,butatealmostnomeatinCase1)andindividualvariationinperceptionsofwhatisediblemayexist.
Alargeproportionofscavengingcaseshaveoccurredwhenchimpanzeesreturnedtositesofrecentmonkeyhuntstoeatmeatleftthere(Table1).
Eventhisisrare,probablybecausechimpanzeescompletelyconsumemostpreyimmediatelyaf-terhunts.
AtNgogo,individualsmaykeepmeatovernightaf-terhuntslateintheday,butrarelyleavelargeamountsofmeatatornearkillsites.
Case2wastheonlycaseofscavengingfromtheirownkills,andthemanypotentialconsumersatthemeat-eatingsessionwouldalmostcertainlyhaveeatenev-erythingedibleifnotforthedistractionbyneighbors.
OR'sdiscoveryofacarcassfarfromthehuntsitethefollowingmorningwasfortuitous.
GiventhatOldowanhomininsfacedcompetitionforfreshcarcasseswithmultiplelargeand130D.
P.
Watts/JournalofHumanEvolution54(2008)125e133dangerouscarnivorespecies(VanValkenburgh,2001;Plum-mer,2004),theirforagingtacticspresumablydidnotincluderetrievalofunconsumedmeatfrompreviouskillsites.
Chimpanzeesmayalsobelessinclinedtoexploitscaveng-ingopportunitiesthanarecarnivorousmammalsbecausetheylackcomparablephysiologicalbarrierstoinfectiousagentsincarrion(HamiltonandBusse,1978;cf.
PlummerandStanford,2000;Plummer,2004).
Inparticular,theymayfacehighrisksofcontractingzoonosesfromdeadordyingheterospecicsthattheyhavenotseenpredatorsattack.
Forexample,bothduikerandchimpanzeesaresusceptibletoEbola(Roquetetal.
,2005),andanthrax,botulism,andotherdiseasesaredi-rectlytransmissiblefromcarcasses(HamiltonandBusse,1978).
UseofleopardkillsatMahalesuggeststhatscavengingmightpreviouslyhavebeenmorecommoninhabitatsfromwhichleopardshavebeeneliminated(e.
g.
,Kibale).
However,Mahalechimpanzeesrespondedfearfullytothefreshcarcassofanadultbushpigkilledbyaleopardandthusignoredameatpackagefarlargerthantheirtypicalprey(Nishida,1994).
Additionally,researchershavenotobservedchimpan-zeesscavengingonleopardkillsatTa(whereleopardspreyonchimpanzees;Boesch,1991),althoughthismaybepartlybecauseTachimpanzeesdonothuntduiker,whicharethemajorpreyofleopards(BoeschandBoesch-Achermann,2000;ZuhberbuhlerandJenny,2002).
DosocialfactorscontributetotherarityofscavengingbychimpanzeesHuntingandespeciallymeateatinghavesocialimportance.
Allowingotherstohavemeatmaynotbecostlytohunterswhentheycapturepreytoolargeforasingleindivid-ualtoeat(e.
g.
,anadultmaleredcolobusmonkey)ortootherswhoobtainlargesharesofsuchkills,andcostsofharassmentbybeggarsmaysometimesoutweighthecostsofallowingthemtohavesomemeat(Gilby,2006).
Still,delayedreciproc-ityinmeatsharing,activesharing,andpreferentialsharingbe-tweenmaleallies(MitaniandWatts,2001,unpublisheddata)pointtowardsmeatsharingasameanstomaintainandstrengthensocialbondsbetweenmalesandtodevelopandmaintainal-liances.
Yetinsofarasmeatsharinghasthisfunction,itsef-fectsshouldnotdependonwhetherinitialmeatpossessorsobtainmeatviahuntingorscavenging.
Huntingbymalechimpanzeesmayinvolvecostlysignaling(HawkesandBliegeBird,2002).
Whilemeatsharingdoesnotincreasemalematingsuccess(MitaniandWatts,2001;Gilby,2006),huntingmonkeysisrisky,andmotivationtohuntandskillatpreycapturemaydisplaymalequality(HawkesandBliegeBird,2002)andsignalwillingnesstotakerisksandtoprovidetnessbenetstoothermales.
Suchsignalscouldinuencedecisionsaboutwhentoseekopportu-nitiestoengageinintergroupaggression(WattsandMitani,2001)andcouldinuencealliancestrategies(MitaniandWatts,2001).
Acquiringmeatthroughscavengingwouldnotprovidecomparablesignals.
Costly-signalingtheorymayindeedexplaincertainaspectsofhuntingandfoodsharingbyhumanhunter-gatherers,suchasHadzamen'sspecializationonhuntinglargepreythatbecomepublicgoods(HawkesandBliegeBird,2002)andturtlehuntingbyMeriammenwhoinvestconsiderabletimeandenergyinriskyhuntsbutdonatetheircatchtoothers(BliegeBirdetal.
,2001;Smithetal.
,2003).
However,theultimateexplanationforhuntingbychim-panzeesispresumablythatmeatisanextremelyvaluablenutritionalresource(MitaniandWatts,2001);thisremainstrueregardlessofitssource,solongaseatingitdoesnotimposetnesscosts(e.
g.
,becauseofdiseasetransmission).
Likewise,costlysignalingisrelevanttohuntingbyhumanspreciselybecausemeathassuchhighnutritionalvalue.
Hadzamenspecializeinhuntinglargegamedespitetheabsenceofdisproportionatenutritionalbenetstothemselvesandtheirfamilieswhentheysucceed,buttheyalsotypicallyexploitscavengingopportunities.
Confrontationalscavengingmayprovidesomescopeforcostlysignaling,butitsmainimportanceispresumablynutritional.
Itseemsunlikelythatchimpanzeeswouldfailtopursuesimilarnutritionalbenetssimplybecausepassivescavenginglackspotentialforcostlysignaling.
AtNgogo,chimpanzeescaptureonlyinfantandjuvenileredduikerandthelargestprey(adultmaleredcolobusandblack-and-whitecolobus)arefarsmallerthanthemodalsizeofungu-latespeciesusedbyBed1Olduvaihominins(Plummer,2004).
Scavengingcanincreasethesizerangeofcarcassesfromwhichchimpanzeesgainmeat(Table1):theadultredduikerinCase1waslargerthanallpreyitems(althoughRAandBIatealmostnoneofit),asweretheadultbushbuckscavengedatMahale(Hasegawaetal.
,1983).
However,itistooraretoproviderealnutritionalbenet.
Incontrast,scavengingcarcassesofmedium-sizedbovidsandlargermammalsprobablyaffordedanutritionallysignicantamountofmeattoOldowanhominins(PlummerandStanford,2000;Bunn,2001;Dominguez-RodrigoandPickering,2003;Plummer,2004).
Scavengingmighthavebeenmorecommoninmiombowoodlandhabitatsattheextremeofchimpanzeeecologicaltolerances(e.
g.
,Ugalla)thaninrainforestsandmorehumidwoodlands,butrecenthuman-inducedchangesinpredatorguildsmakeinvestigatingthispossibilitydifcult(McGrew,1992;Moore,1996).
LatePlioceneandearlyPleistocenehom-ininhabitatsexperiencedagradualtrendtowardsgreateropen-ness,withincreasedgrasslandafterabout2Ma,althoughvariationwasapparentlyextensive,withopengrasslands,bushlands,locallyforestedareas,andshrublandswithwet-landsallreconstructedforlocalitiesatwhichfossilsofHomoand/orParanthropushavebeenfound(ReedandRec-tor,2006).
Suchvariation,plusvariationinthelocaldensities,foragingstrategies,morphology,andgregariousnessoflargecarnivores,wouldhaveledtowidevariationinthefrequencyofscavengingopportunitiesforOldowanhominins(Plummer,2004),butopportunitiesshouldhavebeenmorecommonthanforchimpanzeesinforesthabitats.
TherecentdiscoveryofmiddlePleistocenechimpanzeefossils,datedtobetween5453kaand28412ka,intheKapthurinFormationoftheTugenHills,Kenya(McBreartyandJablonski,2005),castsdoubtonthenotionthatchimpanzeeevolutionaryhistoryhasbeendevoidofopportunitiestoscavengesimilartothoseavailabletoearlyHomoandofexposuretothesamepredatorguilds.
Thesefossilsshowthatthegeographicrangeof131D.
P.
Watts/JournalofHumanEvolution54(2008)125e133chimpanzeespreviouslyextendedintotheEastAfricanRiftValleywheretheywerepotentiallysympatricwithHomoinsemiarid,butlargelywooded,habitats(McBreartyandJablon-ski,2005).
Nevertheless,scavengingwouldprobablynothavebeenamajorcomponentofchimpanzeeforagingstrategiesinsuchacontextforseveralreasons.
Contemporaneoushomininsprobablycoulddrivelargecarnivoresfromkillsbythrowingrocksforcefullyandaccurately(VanValkenburgh,2001);chimpanzeeslackthisability,andtheydonotusetoolstohunt.
Additionally,wildchimpanzeeshaveneverbeenseenproducingsharp-edgedstoneakesofthesortthatwouldhaveallowedearlyhomininstoremovemeatfromcarcassesquickly,thenleaveacquisitionsitesandtherebyreducetheirownriskofpredation(Foley,1987;VanValkenburgh,2001);ancestralchimpanzeespresumablyalsolackedthisability.
Chimpanzeescanopenthelongbonesofcolobus-sizedpreywiththeirmolarstoextractmarrow(Teleki,1975;personalobservation),buthaveneverbeenseenusinghammerstonestoopenbones;withoutthisability,theycouldnothaveex-tractedmarrowfromlargercarcassestothesameextentasdidOldowanhominins.
Chimpanzeesinsemiarid,locallywoodedhabitatintheMt.
AssirikregionofSenegalareatriskofpredationbylionsandhyenasandarenotknowntoscavengefromeitherofthesecarnivores;thesamemightgenerallyhavebeentrueof''savanna''chimpanzeesincomparable,relativelyopenhabitats(Moore,1996;HuntandMcGrew,2002).
Stanford(2001:129)statedthat''meateatingpatternsbywildchimpanzeeslendlittlesupportforhypothesesforscav-engingasamajorcomponentoftheearlyhumandiet.
''Thisobservationmightberephrasedtostatethatdataonchimpan-zeecarnivoryhavelittlerelevancetohypothesesabouttheim-portanceofscavengingoncehomininsstartedtoincludemeatfrommedium-to-largemammalsintheirdiets,otherthantohighlightquestionsaboutdifferencesinhabitatpreferencesandotheraspectsofbehavioralecology,morphology,andcog-nition.
PassivescavengingmighthaveofferedlittlerewardtoOldowanhominins(Blumenschine,1991;Tappen,2001),anddeliberatesearchforconfrontationalscavengingopportunitiesmighthavebeenenergeticallyinefcientandrisky(Tappen,2001).
Nevertheless,iftheforagingstrategiesofsomePlio-cenehomininsincludedenoughscavengingtoyieldimportantnutritionalgains,andiftheycommonlyengagedinconfronta-tionalscavengingfromlargecarnivores,thenthesebehaviorswouldhavedistinguishedtheirbehavioralecologyfromthatofchimpanzees(PlummerandStanford,2000;Bunn,2001;Schoeningeretal.
,2001;Plummer,2004).
AcknowledgementsIthanktheMakerereUniversityInstitutefortheEnviron-mentandNaturalResources,TheUgandaWildlifeAuthority,andtheUgandaNationalCouncilforScienceandTechnologyforpermissiontoworkatNgogo.
IamindebtedtoJohnKase-neneandGilIsibirye-BasutaforpermissiontoworkattheMakerereUniversityBiologicalFieldStationandforalltheirsupportovertheyears.
JohnMitaniandJerryLwangahavebeenoutstandingcolleagues,collaborators,andfriendsatNgogoandhavecontributedgreatlytomyunderstandingofhuntingandmeatsharingbychimpanzeesthere.
FieldworkatNgogoispossibleonlywiththeexpertassistanceofGod-freyMbabazi,AdolphMagoba,LawrenceNdagezi,andAlfredTumusiime.
MyresearchatNgogohasbeensupportedbytheL.
S.
B.
LeakeyFoundation,theNationalGeographicSociety,PrimateConservation,Inc.
,andYaleUniversity.
BillKimbel,ManuelDominguez-Rodrigo,andananonymousreviewerpro-videdextremelyhelpfulcommentsonanearlierversionofthismanuscript.
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