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The Food-SharingBehavior of ProtohumanHominids

by Glynn Isaac April 7978 Excavotions ot two-million-yeor-oid sites in Eost Africo ofler new insightsinfo humon evoiutionory progrss by showing thot eorly erect-stonding hominids mode tools ond coried food to q ftome bose

patthe pastdecade investigators these observationscan now be corn- lacts:for example the distribution /a\ver ot fossil man pared quantitative have discovered with data from antern of the discarded tools in different f I \-z the remainsof many ancient othernewareaofstudy,namely thecul settings and the association of toolswith protohumansin East Africa. Findings tural ecologyof human societies that various kindsof food refuse. A studyof at Olduvai. Laetolil. Koobi Fora. the support themselveswithout raising the contextsof the early African artiOmo Valley and Hadar, to name some plants or animals:the few surviving factsyieldsuniquecluesboth to the ecoprominent locations,make it clear that hunter-gatherers of loday.Anotherim- logical circumstances of the protohubetween two and three million years portant new movementhasinvolvedthe man toolmakersand to aspects of ther agoa numberof two-legged hominids, es- direct study of lhe ecological circum- socioeconomic organization. sentially humanin form, inhabiled lhis stances surrounding human evolutionpart of Africa. The paleontologists who ary developments. Investigations ol rhis Comparing Men and Apes have unearthedthe fossils report that kind have becomepossiblebecause the they differ from modern mankind pn- stratifiedsedimentary What are the patterrsof behaviorthat rocks of East Af' marily in being small. in having rela- ricapreserve. in addition to fos:ilhomi set the species aparl from Hono sapiens tively large jaws and teeth and in hav- nid remains, an invaluable store ofdata: its closest living primaterelatives? Il is ing brainsthat, althoughthey are larger a coherent, ordered recordof the envi- not hard to draw up a list of suchdifferthan thoseof apesof comparablebody ronments inhabited by these protohu- ences by comparing humanandapebesize,are rarely more than half the size mans. havior and focusingattentionnot on the of modern man s. The work of the archaeologist rn many featuresthe two have in common The African discoveries have many drawing inferencesfrom such data rs but on the contrastingfeatures.In the implicationsfor the student of human made possibleby the fact that at a cer- list that follows I have drawn on recent evolution.For example, one wondersto tain stagein evolution the ancestors of field studiesof the great apes(particuwhat extent the advancedhominids of modern man becamemakersand users larly the chimpanzee,Pan troglodytes) two million years agowere human in of equipment. Amongotherthings. they and on similarstudies of the organizatheir behavior.Which of modernman s shaped, usedand discarded numerous tion of living hunter-gatherer societies. special capabilitiesdid they share? stone tools. These virtually indestruc- The list tendsto emphasize the contrasts what pressures of naturalselection, in tible artifacts form a kind of fossil rec- relating to the primary subsistence adthe time sincethey lived. led to the evo- ord of aspects that is, the questfor food. of behavior.a record aptation, lutionary elaborationof mans mind that is complementary to the anatomr' First. Hotno sapiens 1s a twolegged and culture? These are questionsthat cal record provided by the fossil bones primate who in moving from place to paleontologists find dimcult to answer of the toolmakers themselves. Students placehabitually carries tools,food and because the evidence that bearson them of the Old StoDeAge once concentrat- otherpossessions with hisarmsor either is not anatomical. Archaeologists,by ed almostexclusively on what could be in containers.This is not true of the virtue of their experiencein studying learned To- great apeswith regardto either posture from theform of suchtools. prehistoricbehaviorpatternsin general, day the emphasis in archaeology is in- or possessions. can help to supply the answers. creasingly on the contextof the artiSecond. members of Hono sapiens soIt has long beenrealizedthat the human species is set apart from its closest Iiving primaterelativesfar more by dif- pAsr AND PRESENT LANDscApES in rhRift valteyregion of East Africa, shown scbcferences in behavior than by differences in anatomy. Paradoxically, however, the study of human evolution has traditionally been dominated by work on the skeletal and comparative anatomy of fossil primates. Seviral new researci mouements in recent years. to*eu"r. nave oegun to oroaoen rne scope oI direct evolutionary inquiry. One suctr movement involves investigations of the behavior and ecology of living primates and of other mammals. The results of page, prserveit maticdty on the opposite summari? th geological ;cti;ity rhat 61gt anit tatel exposed evidence oi protohurnan tife. Two mi ion ycarsago (trl) the bonei of hominids(1-.1, crlrr) and other animals(,rJ, rrlor) weredistributed across hills snd a floodplain(loregmuno rdiacnt to a Rift vallcy laka Also lying on th surfacewr stonc tools (r/ark do.r) msd' lnd tools uscd.nd discarded by th protohumens, Layrsof sedimcnts then covered tb bones lying on-_rhc flood_plain; burial presrvd them,wheieas thc boncsand tools in th hills wd afrer s fault hasraiseda block or sediments, elosion ."""nlu1lll :1:l"9"1yilll'1i1fto4,,,). is exposinq somof lhc lone-buried bones and clunersof lools.includingthe lhreetypesofsile .loJnonitr".u*u""inthe-lopblockdiagram{,4-c},sitesofTypeAconlainclustersofstone ,h" ;i.;; .i;;; corsthat providdihe law marerialror rhe toots and *1.1.r"1"f;tii process, wastef;kcs from rhe roolmaking but tittfi or no boneis present. sitesof TypGB contain simitarclustcrs of toolsin asiociation with the bones of a singielargeanimal.Siiesof Type C aho coniainsimilar cluslers of tools.but the bones ar from manv difrrcntanimalsDcies.


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HUMAN ANCESTORS home base is evident in the social arrangements of the great apes. Fifth, humanhunter'gatherers tendto devote more lime thanotherliving primatesto the acquisitionof high-protem foodstutrsby hunting or frshingfor ammal prey. It should be noted that the distinctionis one not of kind but of degree. Mountirg evidenceof predatory behavioramonggreat apesand monkeyssuggests that the principal contrast between human beingsand other living primateswith respectro predarionis that only human beingshabitually feed on prey weighing more than about 15 kilogams. The gathering activities of human hunter-gatherers include the collection of edibleplants and small items of anlmal food (for examplelizards. turtles, frogs,nestlingbtds and eggs). Characteristically a proportion of thesefoodstuffs is not consumeduntil the return to the home base.This behavior is rn marked contrast to what is observed among foraging great apes, which almost invariably feed at the spot where the food is acquired. Still another conrast with great-ape feedingbehavioris human hunter-gatherels'practiceof subiecting many foodstuffs to preparationfor consumption, by crushing,grinding,cutting and heating. Such practices are not observed among the great apes. Human hunter-gatherers also make useof variouskindsof equipmeotin th questfor food. The human societywith pelhapsthe simplest equipment everobserved wasthe aboriginalsocietyofTasmania,a populationof hunter-gatherers that wasexterminated io the 19thcentury. The inventory of the Tasmanian$' equipmentincludedwood clubs,spears and diggingsticks,cuttingtools madeof chipped stone that were used to shape the wood objects,and a variety of containers: trays, basketsand bags. The Tasmanians also had 6re. Although such equipmentis simple by our standards, it is fa! more complex than the kind of rudimentary tools that we now know living chimpanzees may collect and use in the wild, for exampletwigs ano grasssrems. ln addition to this lengthylist of subsistence-related behavioral contrasts betweenhuman hunter-gatherers and living primatesthere is an entire realm of other contrasts with respect to socialorganization. Although these impoltant additional featuresfall largely outside the range of evidenceto be considered here, they are vital in defining human patterns of behavior.Among themis the propellsity for the formation of long-

cietiescommunicateby meansof spoken language; suchverbal communication serves for the exchange of information about the past and the future and also for the regulationof many aspects of social relations.Apes communicate but they do not have language. 'flwd, n Homo sapr'ens societies the acquisition of food is a corporate responsibility, at least in part. Among membersof human socialgoupings of various sizesthe active sharingof food is a characteristicfolm of behavior: most commonly family groups are the crucial nodesin a network of food exchange. Food is exchangedbetween adulti, and it is sharedbetweenadults andjuveniles. The only similar behavior observed arnong the great apes is seen when chimpa[zees occasionally feed on meat.The chimpanzees' behavior,howeyer, falls far short of active sharing;I suggest it might bette!betermedtolelated scrounging. Vegetablefoods, !r'hich are the greatapes'principal diet,are not shared ald are almost invariably consumed by each individual oD the spot. Fou h, in human social groupings there exists at any given time what can be called a focus in space,or "home base," such that individuals can move independently over the sunoundiogterrain and yet join up again. No such

DESOLITE LANDSCAPE in the alid Koobi Forr district of Kenya b typicrl of thc ktnd ot erodd trrrin where gullying erposesboth boDcs and stonc tools lbrl wer burid bencatbsedhhents anil volcanic ash E|or lhao r million ye$s rgo. Excrvrtlon ln progre6s(cater)

16xposiDg lh bippopotrmus bon6.rndclustersof rrtlf.cts th.t hart been prrtially bared by recnt erosion and were founil by Rich$d Leaky in 1969, The site ls typlcrl ol the kind thrt lncluils the ro. m.ins of a singl anim.l rnd m.ny tools mrnuhclured on ihe spot


term mating bondsbetween a male and oneor more females. The bondswe call "marriage involvereciprocal econom. ic ties, ioint responsibilityfor aspects of child-rearingand restrictionson sexual access. Another suchsocialcontrast iE evident in the distinctively human propensity to categorize fellow members of a group according to kinship and metaphorsof kinship. Human beings regulate many social relations. mating included.accordingto complex rulesinvolvingkinshipcategories. Perhaps family ties of a kind exist among apes, but explicitcategories andrulesdo not. These differences are emphasized by the virtual absencefrom observed ape behavior of thosedistinctivelyhuman activities that arecategor izedsomewhatvaguely as"symbolic and ritual.' Listing the contrastsbetweenhuman and nonhuman subsistence strategies is inevitablyan exercise in oversimplification. As has beenshown by contemporary field studiesof various great apes and of human beingswho. like the San (formerly miscalled Bushmen) of the Kalahari Desert, still support themselveswithout farming, there is a far greaterdegree of similarity between the two subsistence strategies than had previously beenrecognized. For example, with regardto the behavioral repertories involving meafeating and tool-using the differences apeand man are between differencesof degree rather than of kind. some scholars have even usedthe data to deny the existence of any fundamental differnces betweenthe human strategies and the nonhumanones. It is my view that significanr diFerences remain. Let me cite what seemto me to be the two most important. First, whereas humansmay ieed as they fordo not regularly agejust asapes do, apes postponefood-consumptionuntil they havereturnedto a home base. ashuman beings do. Second,human beings actively sharesome of the food they acquire.Apesdo not, eventhoughchimpanzees of theGombeNationalParkin Tanzaniahavebeenobserved to tolerate scrounging when meat is availableFrom Hominid to Human


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PROMINENT SITES in East Alricr include (fiorh nortb to south) Hailar, Mlkr Kuntur6 and Shungun in Ethiopia, the Koobi Fora ilistrict to tbe sst of Lrke Turkana in Kenya, Chesowanja in Knya and Pnfui, Olduvri Gorg and Laetolil in Tananir, Dates foi clustrs of 6tone lools, some associated with animrl bons, uncoverd rt lh6e sites rangc from on hillion yerlr ago (Olduvri Upper Bed lI) to 2.5 nilliotr (Had.r uppr beds). Some sites mry bc e}.en older.

UPPER BEDII M I D O L E E Dl l 1.4 LO$rEnaEO tl





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1_6 BEOI 1.8 2.O 2.2 2.4 KAS,HAS


? Two complementary puzzles face HADABUPPER BEDS 2.6 anyonewho undertakes to examinethe questionof human origins.The fust re- (D 2.8 latesto evolutionarydivergence. When did the primate stock ancestralto the ut 3.0 living apesdivergefrom the stockanto man? What were cestral the circum3.2 LOWEF BEDS HADAB stancesof the divergence? Over what geographical rangedid it take place?It is not yet establishedbeyond doubt LAEfOLIL whetherthe divergence occurreda mere five to six million Vincent RELATM of selected sites in Eest AIiic! is indiceted in this table. Olduvai M. Sarichof the University of Califor- Gorge bds I ANTIQUITY anrt ll range fron l.E to 1.0 nillion years in agc. Tbe Sbungura sites in tbe Omo nia at Berkeleyand othersargueon bio- Vdly are more thrn two million yeals old. Two Koobi Fora locsles, tbe hippopotamus/artichemical grounds,or 15 to 20 million f.ct site (HAS) and th Kay Behrensmycr site (KBS), are at least 1.6 hillioo yeais old, Initial years ago, as many paleontologists be- geologicd s,tudie6of th Koobi Forr sites suggestdthrt tbey mi8ht be 2.5 million years olrt lieve on the groundsof fossil evidence. (coloredline\, Only hominid fossils have ben found ir the lower bds at Hadr.nd ai Laetolil.


HUMAN ANCESTORS The archaeological research that has inspiredthe formulation of newhypotheses concerning human evolutionbegan nearly 20 yearsago when Mary Leakey and her husbandLouis discoveredthe fossilskull he named"zinjanthropus"at Olduvai Gorge in Tanzania.The excavations the Leakeys undertook at the site showed not only that stone tools werepresentin the samestratathat held this fossil and other hominid fossilsbut alsothat the discarded artifactswereaswith numerous anisociated broken.up mal bones.The Leakeystermed these 'livconcentrations of tools and bones ing sites." The work has continued at Olduvaiunder Mary Leakey's direction, and in 1971 a major monograph was publishedthat hasmadethe Olduvai resultsavailablefor comparativestudies. Other important opportunities for archaeological research of this kind have cometo light in the GregoryRiftValley, at placessuch as the Koobi Fora (formerly East Rudolf) region of northem Kenya, at Shungarain the Omo Valley of southwestern Ethiopia and in the Hadar regionof easternEthiopia. Current estimates ofthe ageofthesesitescovera span of time from about 3.2 million yearsago to about 1.2 million. with Since1970I havebeenco-leader Richard Leakey (the son of Mary and Louis Leakey) of a team working at Koobi Fora. a district that includesthe northeasternshore of Lake Turkana (the former Lake Rudo10.Our research on the geology, paleontologyand paleoaothropology of the district involves the collaborationof colleagues from the National Museum of Ke[ya and from many other parts of the world. Work begannin 1968and hashad the help and encouragement of the Covemment oi Kenya, the National ScienceFoundationandtheNational Ceographic Society. Our investigations have yieldedarchaeological evidence that corroborates and complementsthe earlier evidence from Olduvai Gorge. The combined data make it possibleto se iust how helpful archaeology canbe in answering

At leastonefact is clear.The divergence took placelong before the period when the oldest archaeological remains thus far discovered first appear.Archaeology, at leastfor the present, can make no contribution toward solviog the puzzle of the split betweenancestralape and ancestral man. puzzle, As for the second fossilevidencefrom East Africa showsthat the divergence, regardless of when it took place,hadgivenrisetwo to threemillion years ago to populations ol smallish twoleggedhominids.The puzzleis how to identify the patternsof natulal selection that transformed these protohumans into humans.Archaeologyhas a majorcontriburion to makein elucidating the secondpuzzle. Excavation of protohuman these sites hasrevealed evidencesuggesting that two million years ago someelements that now distinguish man from apeswerealreadypart of a noveladaptivestrategy. The indications are that a particularly importantpart of that strategywas food-sharing.


KOOBI FORA ARTIFACTS includfoulfrom the HAS assemblage (lel0 a four frorn the KBS esshblege (riano. AII are shown actual siz; the slone is besalt Tbe HAS cor (d) shows what is ldt of a piece of stone alter a number of fak5 have been struck ftom it by prcussion. Ibc jrg8ed edges produced by fake iehoval give the cor po.

tential usefulness as a tool. The frkes were iletrched frorr the core by blows with a brmmelstone like tbe one shown here (c). The shary edgesof tbe flakes,sucb as lbe example illustrated (r), allow tbeir use ss cutting tools. The tiny flake (d) is probably an accidentrl proaluct of lhe percussion procss; the presnce of many stonc splinters such



questions concerning humanevolution. At Koobi Fora.asat all the other East African sites,depositsof layered sedi, which accumulared ments, long ago in the basinsof Rift Valley lakes.are now beingerodedby desert rainstorms and traDsientstreams.As the sedimentary bedserode,a sampleof the ancientarti, factsand fossil bones they containis exposed at the surface.For a while the exposed material lies on the ground. Eventually, however, the fossil bones are destroyed by weathering or a storm washes away stoneand bone alike. All field reconnaissance in East A1nca progresses along essentiallysimilar lines. The field teams search through erodedterrain looking for exposed fossils and artifacts.In placeswhere concentrations of fossil bone or promising archaeological indications appear on the surfacethe next step is excavation. The digging is done in part to ulcoverYurther specimens that are still in place rn the layers of sedimentsand in part to exactinformalion abourrheorigSalher

inal stratigraphic locationof the surface material. Most important of all, excavalion allowsthe investigalors to plot in detail the relativelocationsof the material that is unearthed.For example,if thereare associations amongboDes and betweenbones and stones.excavation will revealthesecharacteristics of the slte. The Types of Sites The archaeological tracesof protohuman life uncovered in this way may exhibit severaldifferentconfigurations. In someancieDt layerswe havefound scatteringsof sharp-edged broken stones evenrhough lhereareno otherstones in the sediments. The brokenstones come in a rangeof forms but all are olthe kind producedby deliberate percussion, so that we can classify them as undoubt, ed artifacts. Suchscatterings of artifacts areoften found without bonebeingpresent in signincant amouots. These I proposeto designate sites of Type A.

ln someinstances a layer of sediment may include both artifacts and animal bones. Suchbone-and-artifact occurrences fall intolwo categories. I hefirstconsistsof artifacts associated with bones that representthe carcassof a single large animal: these sites are designated Type B. The secondconsistsof artifacts associated with bonesrepresenting the remains of several different animal species; thesesites are designated Type C. The discoveryof siteswith thesevaried configurationsin the sedimentsat Koobi Fora and Olduvaiprovides evrdence lhat whenthe sediments containingthemwerebeing deposited some2.5 to 1.5 million yearsago, therewas at least onekind of hominidin EastAfrica that habitually carried objects such as stonesfrom ooe place to another and madesharp-edged tools by deliberately fracturing the stonesit carried with it. How doesthis archaeological evidence match up with the hominid fossil record? The fossilevidence indicates that



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as this on in rh HAS tool clustls indicates thatthe stone tools w.e made on th spot At the same timc the absenc of tocat unworkcd stone as potential raw mateiial for tools suggeststhat the cores were canied to tb site by the toolmakers. The artifacts from the second assemblagealso include a core () that bas had many flakes removed

by percussion and anothersmall splinter of stone (t). The edgesof th two flakes (l a)are sharp cnougb lo cut mal, hide, sinew or wood. As al the hippopotamus/artifact site, tte absence of local raw materi.l for stone tools at tb Kay Behrenshyer sit suggeststbat suitabl lumps ot lava must hsve ben transportd there by the toolmakN,

HUMAN ANCESTORS of future investigators. Here I shall simply discuss what we can discoverabout lhe activities of earlytoolmaking hominidswithoutartempting to identify their taxonomicposition (or positions). Readingthe Evidence As examples oi the archaeological evidenceindicativeof early hominid paf ternsof subsistence and behavior,consider our findings at two Koobi Fora excavations. The fust is a locality catalogued as the hippopotamus/artifact site (HAS) because of the presence of fossilized hippopotamus bons and stonetools. The site is 15 miles eastof Lake Turkana. There in 1969 Richard Leakey discovered an erosiongully cutting into an ancientlayer of volcanicash known as the KBS tuff. (KBS standsfor Kay Behrensmeyer site: she, the geologistpaleoecologist of our Koobi Fora reKOOBI FORA LANDSCAPE in ihc vicinity of the hippopotaDus/rrtitact site consisted of r search team,first identifiedthe ashlayer level foodpkin ngr tbe mergin of a leke (ro? s?.tior). Protobumrn foragers rpparently foutrd at a nearby outcrop.) The ash layer ls tbe c$cass of. hippopotamus lying in a strerm-bed bollorv and made tools on tbe spot in odr to butcher ihe carcass. Their rctions left a scatter of stone tools among the bones .nd on ih part of a sedimentary the uppermost deground neerby, The floodplain w.s bu d ulder layers of silt end ash and was sub6quently posit known to geologists as the Lower crodeil lbotton yction), exposing som bones and tools. Tbir discovery led to excsvation. Member ol the Koobi Fora Formation; herethe ashhad filled in oneof the many dry channels ofan ancientdelta.Leakey two andperhaps threespecies of bipedal man weremaking the stonetools.These found many bonesof a singlehippopothominidsinhabitedthe areaat this time, are the fossilforms, of early Pleistocene amus calcass weathering out of the so that the questionarises: Can the spe- age, classifiedby most paleontologists erodedash surface,and stone artifacts cies responsible for the archaeological as an early species of the Eenus Homo. lay amongthe bones. evidence be identified? The questionof whetheror not contemJ. w. K- Harris, J. Onyango-Abuje For the momentthe bestworking hy- poraneous hominid species of the genus and I supervised an excavationthat cut pothesis seems to be that thosehominids Australopithecus alsomadetoolsmust be into an outcropwherethe adtacent delta that were directly ancestralto modern setasideas a challenge to the ingenuity sediments had not yet bendisturbedby erosion.Our digging revealedthat the hippopotamus carcass had originally lain in a depressioror puddle within an ancientdeita channel.Among the hippopotamus bones and ir the adjacent steam bank v/e recovered119 chipped stones:most of them were small sharp flakesthat, when they are held between the thumb and the frngers, make effective cuiting implements. We also rcoveredchunksof stonen ith scarsshowiog that flakeshad been struck from them by percussion. In Paleolitiic tool classification theselarger stonesfall into the categoryof core tool or chopper.In addition our digging exposeda rounded river pebble that was batteredat both ends: evidently it had been used as a hammer to strike flakesfrom the stone cores. The sediments where we found these artifactscontainno stones larger than a pea.Thus it seems clear that the makers of the tools had carriedthe stoneshere irom somewhere el$e.The association betweenth patch of artifacts and the hippopotamus bones further suggests that toolmakerscame to the site carrying stonesand hammeredoff the small sharp-edged ffakeson the spot in order to cut meatfrom the hippopotamus carIIAMMERSTONE unartbed at the bippopot mus/artifact site is . six.centirnetr basrlt pebcass. We haveno way of telling at presbl; it is shown her being lilted from its position on lh sncient ground surface sdi.cnt to the hippopotrmus bones. Worn srnooth by r.l.r ent whetherthe toolmakersthemselves .ction before it caught ihe eye of . toolmskcr soln 1.7 orillion yerrs .go, the pebble is bltter.d .t bolb ends ss a result of use as a hrrnrnr. killed the animal or only came on it



dead.Given the low level of stonetechI am inclinedto susnology in evidence, rather than hunting pect scavenging The HAS depositwas formed at least 1.6 million yearsago.The archaeologlthat the be cal evidencedemoDstrates havior of some hominids at that time differed from the behavior of modern

great apes in lhaL these prolohumans not only made cutting lools but also ate of large ani meal from the carcasses mals. The hippopotamus/artifact site thus provides cortoboration for evtdence of similar behavior just as long ago obtained from Mary Leakey's excavations at Olduvai Gorge.

This finding does not answerall our questions. Were these protohumans roaming the landscape,foraging and hunting. in the way that a troop of baboons does today? Were they instead hunting like a pride of lions? Or did someother behavioralpattern prevail? Excavation of another bone-and-arti-




FINDINGS at the hipFoPot.mus/artifactsite are shown schematically in this block diagram; squ.res are one mter io a side. ln the fore;rountl sre lbe obiects lhar had bcen exPosedbv weathering: bippoootsmu! limb bones (a-d) and reerh t'mall oper\). man) frrgments of bonc (sror"/ ddtrss) and a few stone artitaets (colorcd dots) Trenching (dasn.d ti e, color' ^.rlt billside excavation over a wid

a.ea exposed an ancient soil surface (drlor) overlying a deposit of sil(ircler' tr iufi, Lvine on lbe ancicnl surface $ere slone cores loper strarp'edgea flakes had ben \trucl' mor then i"tort t,i^ "t'icl 100 other stone arlifacts and more than 60 additional fragmnts of teth and boncs. The scatter of tools 5nd broken bones suggeststhe hypothesis thal the toolmakeN fed on meat from the hippoPotamus'


HUMAN ANCESTORS giraffe,pig. porspecies, hippopotamus, cupine and such bovids as waterbuck, gazelleand what may be either hartebeestor wildebeest. It was this site that was designated KBS. The site obviously represented the secondcategory of bone-and-artifacl associationsr roolsin association with the remains of many differentanimal species. Geological evidence collected by A. K. Behrensmeyer of Yale University and othersshowsthat the KBS deposit h a da c c u m u l a t e od n t h es a n d y b e do f a streamthat formed part of a smalldelta. At the time when the toolmakersused the streambed,waterhad largelyceased to Row. Such a site was probably favored as a focusof hominid activity for a number of reasons. First, as every beachgoer knows. sand is comfortable to sit and lie on. Second, by scooping a hole of Do great depth in the sand of a streambed one can usually find water. Third, the growth of trees and bushes in the sun-parched floodplainsof East Africa is often densest alongwatercourses,so that shadeand plant foods are availablein theselocations.It may also be that the protohumantoolmakers who left their discards heretook shelterfrom predators by climbing trees and also spenttheir nightsprotectedin this way. Much of thisis speculative, of course, but we have positive evidencethat the objectsat the KBS site did accumulate in the shade. The sandysiltsthat cameto cover the discarded implements and fracturedboneswere deposited so gently that chipsof stonesmallenoughto be blown away by lhe wind werenol disturbed.In the samesilts are the impressionsof many treeleaves. The species of treehasnot yet beenformally identifled, but Jan Gilette of the Kenya National Herbarium notes that the impressions closelyresemblethe leavesof African wild fig trees. Carrying Stonesand Meat As at the hippopotamus/artifactsite, we have established the fact that stones

fact association, only a kilometer away from the h ippopotam us/arl ifact site. has allowed us to cary our inquiries further. The secondsite had beenlocatedby Behrensmeyer in 1969. Erosion wasuncoveringartifacts,togetherwith pieces of broken-upbone, at anotheroutcrop of the samevolcanicashlayer that contained the HAS artifacts and bones. With the assistance of John Barthelme of the UniversityofCalifornia at Berkeley and others I beganto excavatethe site. The work soon revealeda scatter of several hundred stone tools in an area 16 metersin diameter.They rested on an ancientground surfacethat had beencoveredby layersof sandand silt. The conceltration of artifacts exactly coincidedwith a scatterof fragmented bones. Enoughof them,teethin particular, were identifiable to demonstrate that partsof theremains of several aniwerepresent. mal species John M. Har ris of the Louis Leakey Memorial Institute in Nairobi recognized, amongother

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BONES AND STONE TOOLS were also found in abundance at the Kay Bebrensmeyrsita As th plot of bonedisbibution (a) shows,tbe animal iemains represent many difieient species.Thes are identificd by capitsl lctters; if the nnd was a tooth the letter is circled. Most

arc small to medium-sized boyids, such as gazelle, waterbuck and hartebee$(A). The remains of crocodile (C),girafie(G), hippopotamus (H), porcupine (P) and extinctspecies of pig (J) wer also present. Dots and dasheslocate unidentified teeth and bone fragmenti respectively.

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ISAAC I THE FOOD.SHARING BEHAVIOR OF PROTOHUMAN HOMINIDS largerthan the sizeof a peado not occur naturallycloserto the Kay Behrensmeyer site than a distance of three kilometers. Thus we know that the stoneswe found at the sitemust havebeencarried at leastthat far. With the help of Frank Fitch and Ron Watkins of the University of London we are searching for the specificsources. It does not seem likelythatall theanimals of the differentspecies represented among the KBS bonescould have been killed in a shortinterval of time at rhis one place.Both considerations encourage the advancement of a tentativehypothesis: Like the stones, the bones were carried in, presumablywhile there was still meat on them. If this hypothesis can be acceptd, the Kay Behrensmeyer site provides very early evidence for the transportof food as a protohuman attribute. Today the carryingof food strikesusasbeingcommonplace,but as SherwoodWashburn of the Universityof California at Berkeley observedsome years ago such an actionwould strikea living apeas being novel and peculiar behavior indeed,In short,if the hypothesis can be accepted, it suggests that by the time the KBS depositwaslaid down various fundamental shifts had begunto lake place in hominid social and ecologicalarrangements. It should be noted that other early sitesin this categoryare known in East Africa. so that the Kay Behrensmeyer site is by no means unique.A numberof suchsiteshave beenexcavated at Olduvai Gorge and reported by Mary Leakey. Of thesethe best preserved is the "Zinjanthropus"site of Olduvai Bed I, which is about 1.7 million years old. patchof discarded Here too a dense arti facts coincideswith a concentration of broken-upbones. There is an even larger number of Type A sites (where concentrations of artifactsare found but bones arevirtually or entirelyabsent). Someareat Koobi Fora: others are in the Omo Valley, whereHarry V. Merrick of Yale Uni-


versity and Jean Chavaillon of the French National Center for Scientific (CNRS)have recentlyuncovResearch eredsitesof this kind in members E and F of the Shungura Formation.The Omo sitesrepresent the oldestsecurelydated artifact concentations so far reported anywherein the world; the tools were depositedsome two million years ago. Oneofthe Olduvaisitesin this category seemsto have been a "factory": a quaffy where chert, an excellent tool material,wasreadily availablelor flakwith ing. The other tool concentrations, very few associated bones or noneat all, may conceivably be interpreted as foci of hominid activity where lor one reason or anotherlarge quantitiesof meat werenot carriedin. Until it is possible to distinguish between sites where bone was never presentand siteswhere the boneshave simply vanished because of such factors as decay, however, these will remain difficultto interdeposits pret in terms of subsistence ecology. lvhat, in summary,do theseEastAf-

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Tbe plot of rnifrct dishibution (r) shows tbat three of fom stone corfs (open circlesr, most wasle stone (stlares) and flaks and fregmnts of flaks (dots) were found in 12 adjacent squsres. AIso found here wrs an unworked stone (,{) tbag Iike thc cores, must bsve been

canied to th site from r distrnc, Plotting of all tools and bons uncarthed at the site wrs not .tternptd. Numbers in grid squares (c) show how many flakes and bits of was(e stone (.r/or) and fragments of bone (r/dck) werc recolded witbout exact plotting in -rch square.


HUMAN ANCESTORS tered from menacingpredators.Others haveproposed thatwhen the early hominids foraged,they left their young behind at "nest" or "den" sites(in the manner of birds, wild dogsand hyenas)and returned to these locales at intervals, bringing food with them to help feed and wean the young. If we look to the recordeddata concerning primitive human societies,a third possibility arises.Among extant and recently extinct primitive human societies the transport of food is associatedwith a divisionof labor. The society is divided by age and sex into classesthat charactelistically makedifferent contlibutions to the total food supply. Onesignificant resultof sucha division is an increase in the varietyof foodstuffs consumedby the group. To generalize on the basis of many different ethlographicreports,the adult femalesof the society contribute the majority of the "gathered"foods;suchfoodsaremainly plant products but may include shellfish, amphibians and small reptiles, eggs, insects and the like. The adult males usually,althoughnot invariably, contribute most of the "hunted" foodstufs: the flesh of mammals, fishes, bilds and soforth. Characteristically the males and females range in separate groups and each sex eventually brings back to a home baseat leastthe surplus ol its foraging. Could thissimplemechanism, a division of the subsistence effort, have initiated food-carying by early hominids? One cannot dismiss out of hand the models that suggestsafety ftom competitorsor the leedingof nestingyoung as the initiating mechanisms for foodcarrying. Nevertheless, neither model seems to me asplausibleas onethat has division of labor as the primary initiating mechanism. Even if no other argument favored the model, we know for a fact that somewherealong the line in the evolution of human behavior two patterns becameestablished: food-sharing and a division of labor. If we include both patterns in our model of early hominid society,we will at leastbe parslmonl0us. Other arguments can be advancedin favor of an early development of a division of labor. For example,the East African evidenceshowsthat the plotohuman toolmakersconsumedmeat from a far greaterrange of species and sizes of animals than are eaten by such living primatesas the chimpanzee and the baboon.Amoog recenthuman huntergatherers the existence of a division of labor seemsclearly related to the femalesbeing encumbered with children, a handicapthat barsthem from hunting or scavenging. activitieslhat requLe speed afoot or long-range mobility. For the protohumanstoo the incorporation of meat in the diet in signifcant quantitiesmay well havebeena key factorin the development not only of a division

mains we have uncoveredreflect social and economicnodesin the lives of the toolmakers who left behind these ancient patchesof litter. Becauseof the evidencesuggestive of the transport of food to certain focal points, the f,rst question that the modelmust confrontis why early hominid socialgoups departed from the norm among living subhuman primates,whosesocialgroupsfeed as they range. To put it alother way, what ecological and evolutionary advantagesare there in postponingsome food consumptionand transportingthe iood? Severalpossibleanswers to this questioo have beenadvanced. For example, Modl Strategies Adrierure Zihlman and Nancy Tannerof Thesearchaeological facts and indi- the University of Califomia at Santa cationsallow the conshuctionof a theo- Cruz suggestthat when the protohuretical model that showshow at least mans acquirededible plants out on the someaspects of early hominid socialex- open grasslands, away from the shelter istence may have beenorganized. Crit- of trees, it would have been advantaical to the validity of the model is the geousfor them to seizethe plant prodinference that the variousclusters of re- uctsquickly andwithdrawtoplacesshel-

rican archaeologicalstudies teach us about the evolution of human behavior? For one thing they provideunambiguousevidence that two million years ago somehominids in this part of Afnca were carrying things around, for example stones. The samehominids were alsomating simple but effective cutting toolsofstone andwereat timesactive in the vicinity of large altimal carcasses, presumablyin order to get meat. The studies stronglysuggest that the hominids carried animal bones (and meat) around and concentrated this Dortable food supply at certain places.

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CLUSTERED MIXTURE of artifacis anil animal bones rt the Kay Behremmeyer site is evi dent when the stone (rrlor) and bone (r/dcf) plots ar superposed. Combinetions of this kind are somtimcs producd by stram actiotr, but sucb is not likely to be the case her, rs is attsted by tbe prservatio[ of laf ihprssions anil olher rsdily washd-sway debris such a! fftr splintrs of slone. It apperrs instad that the prolobumrnE who made and discailed their toots hre rveie rlso rsponsible for the bon rccumulation becauae thcy met hIe to shar their food,


of labor but also oi the organizationof movements arounda home base and the transportand sharingof food. The model I proposefor testingvisualizesfood-sharingasthe behaviorcentral to a novel complex of adaptations that included as critical components gathering hunting and/or scavenging. and carrying.Speaking metaphorically, food-sharingprovides the model with a kind of central platform. The adaptive systemI visualize,however,could only hayefunctionedthrough the useof tools and other equipment. For example, without the aid of a carryingdevice primatessuchasourselves or our ancestors could not have transportedfrom the field to the home base a suflicient amount of plant food to be worth sharing. An object as uncomplicatedas a bark tray would have seryedthe purpose,but somesuchitem of equipment would have been mandatory. In fact, Richard Borshay Lee of the Universrty ofToronto hassuggested that a carrying devicewas the basicinvention that made human evolution possible. What about stone tools? Our ancestors, like ourselves. could probably break up the body of a small animal,as chimpanzees do. with nothingbut their hands and teeth. It is hald to visualize them or us,however.eatingthe meat of an elephant,a hippopotamusor some otherlargemammalwithoutthe aid ofa cutting implement.As the archaeological evidencedemonstrates abundantlv.

the protohumans of East Africa not only knew how to produce such stone flakes by percussion but also found them so usefulthat they carriedthe raw matelials needed to make the implements with them from place to place. Thuswhercas the existence of a carrying devicerequted by the model remains hypotheticalasfar asarchaeological evidenceis concerned. the fact that tools were used and carried about is amply attestedto. In this connection itshould bestressed that the archaeological evidenceis also silent with regard to protohuman consumptionof plant foods. Both the morphology and the patternsof wear observableon hominid teeth suggest such a plant component in the diet, and so doesthe weight of comparativedata on patternsamong living nonsubsistence human primates and among nonfarming human societies.Nevertheless,if positiveevidence is to be found,we shall have to sharpenour ingenuity,perhaps by turning to organicgeochemical analyses. It is clear that aslong aswe do not correctfor the imbalancecreatedby the durability of bone as compared with that of plant residues, studiesof human evoluiion will tend to have a male bias! As far as the model is concernedthe key questionis not whether collectable foods-fruits, nuts, tubers, greensand eyeDinsects-were eaten.It is whether the$eprotohumanscarried such foods about. Lacking any evidence for the

consumptionof plant foods, I shall fall back on the argumentthat the systemI visualizewould have worked bestif the mobile hunter-scavenger contribution ofmeat to the socialgroupwasbalanced by the gatheler-carrier collection of plant foods.lvhat is certain high-grade is that at sometime during the past several million years just such a division of labor came to be a standardkind of behavior among the ancestors of modern man. A final cautionary word about the model: The readermay have notedthat I have beencarelul about the useof the words hunter"and 'hunting. This is we cannot judge how much of because the meat taken by the protohumansof East Africa came from opportunistic scavenging andhow much wasobtained by hunting. It is reasonable to assume that the carcasses of animals killed by carnivores and thoseof animalsthat had otherwisedied or beendisabledwould alwayshaveprovidedactivescavengers a certainamount of meat.For the p!esent it seemslessreasonable to assume that protohumans, armed primitively if at all, would be particularly effective hunters.Attempts are now under way, notably by ElizabethVrba of SouthAfrica, to distinguish between assemblages of bonesattributableto scavenging and assemblages attributableto hunting,but no indings from East Africa are yet available.For the presentI am inclined to acceDtthe verdict of J. Desmond




BEIIAVIOR PATTERNS that difter in ilegree of organizatiotr aie conhssted in lhese disgnms Living grelt ape6, xemplified brc by th chinpanzee, cxhibit bhrvior pattems that becsme imporlant in humsn volution but tbe prtterG (&l) exist largely as isolated cl. mcnts. Hunting occurs otr a small scalc but lerds only to "lolerated scrounging" ralbr than rctive food.sbaringi similarly, fools sre usd but tool us is nol irtegrrted with huntidg or scroutrging. Tbe euthor's model (c?x&r) integrates these tbree behavior prtlems rnil othels into r coherenl structur. Food-shsring is sen ss a cenhrl sbuctural l.

hetrt, incorponting thc provisiotr of both animal and plant fooils, tbe organization of s home bosc rnd e division of labor. Supportiog tb intgrrtd structur is . nccssery infnshuclure of tool and equlpmnt manufrctur; for exrmplc, without devices for csrrying food" stufis thre could not be r divisiotr of orgadzed foodhrring.ln hodern buman societies(riArr) the food-sharing shucture has undergoDe socioecolomic ehboration. Its infr$huctur now incorporates all of tcchnology, and a matcbing superstructure hes arisn to incorportt other lmcntr oI whrt i6 collectivly calld cultur.


HUMAN ANCESTORS volving some division of labor and a degreeof food-sharingmight well have been able to function even if it had communicativeabilities little more advanced than those of living chimpanzees.In such a simple subsistence system, howeYer, any group with members that were able not only to exchangefood but alsoto exchange informationwould have gained a critical selectiveadvantage over all the rest. Such a group's gathererscould report on scavenging or hunting opportunitiesthey had observed, and its hunters could tell the gatherersabout any plant foods they had encountered. By the sametokenthe finadjustment of social relations,always a matter of importance among primates, becomes doubly impoltant in a social system that involvesfood exchange. Language servesin modern human $ocieties not only for the exchangeof information but also as an instrumentfor social adjustmentand even for the exchange of misinfolmation. Food-sharhgand tie kindsof behavior associated with itprobably playedan important part in the developmentof systems of reciprocalsocialobligations we that characterize all human societies know about. Anthropological research showsthat eachhumanbeingin a group is ordinarily linked to many other members ol the group by ties that are both social and economic. The French anthropologistMarcel a classic "The Gift. published essay, in 1925, showed that socialtiesare usuallyreciprocal in the sensethat whereas benefits from a relationshipmay inirially passin only one direction,there is an expectation of a future retum of help in time ol need. The formation and management of such ties calls for an ability to calculatecomplex chainsof contingenciesthat reachfar into the future. After food-sharing had becomea part of protohumanbehaviorthe needfor suchan ability to plan and calculatemust have provided an important part of the biological basisfor the evolutionof the human intellect. The model may also help explain the development of human marriage arrangements.It assumesthat in early protohumanpopulationsthe malesand lemales divided subistencelabor betweenthem so that eachsexwaspreferentially tappinga differentkind of food resourceand then sharingwithin a social group some of what had beenobtained.In such circumstances a mating system that involvedal leastonemale in "family" food procurement on behalfof each child-rearingfemale in the group would have a clear selective advantage over, for example, the chimpanzees' pattern of opportunistic relations betweenthe sexes. I have emphasized food-sharingas a principle that is central to an understandingof human evolution over the

assemblages of stonetools beganto reflecta greater cultural complexityon the part ol their makers.The complexity rs first shown in the imposition of more arbitrary tool forms; these changes were followed by increases in the number of suchforms. There is a marked contrast between the pure opportunismapparent in the shapes of the earlieststonetools and the orderly array of forms that apTools as Testimony pear later in the Old StoneAge when Of course,the adaptivemodel I have each form is represented by numerous advancedhere reflectsonly a working standardized examplesin each assemhypothesis and not establishedlact. blageoftools. The contraststronglysugNevertheless, thereis sumcient evidence gests that the first toolmakerslackedthe in its favor to justify looking further at highly developedmental and cultural its possibleimplicationsfor the course abilitiesof more recenthumans. of human evolution. For example,the The evidenceof the hominid fossils model clearly implies that early tool- and the evidence of the artilacts togethmaking hominidsdisplayedcertainpat- e! suggest that theseearly artisanswere terns of behavior that, among the pat- noDluman hominids. I imagine that il terns of behavior of all primates. we had a time machineand could visit a uniquelycharacterize site our own species placesuchasthe Kay Behrensmeyer we and set it apart from its closestliving at the time of its originaloccupation, relatives, the greatapes. Doesthis mean would find hominidsthat were living m that the toolmaking hominids of 1.5 social groupsmuch like those of other to two million years ago were in fact higher primates.The differences would "human"? be apparentonly after prolongedobserI would surmisethat it doesnot, and I vation. Perhaps at the start of each day have beenat painsto characterize these we would observe a groupsplittingup as East African pioneersas protohumans. some of its memberswent off in one In summarizingthe contrastsbetween directionand sornein another.All these living men and living apes I put high subgroups would very probably feedrnon the list languageand the cultural termittently as they moved about and phenomenathat are dependenton it. encountered ubiquitouslow-gradeplant We have no direct means of learning loods suchas berries,but we might well whetheror not any of theseearly hom- observethat some of the higher-grade inids had language.It is my suspicion, materials-large tubers or the haunch however, that the principalevolutionary of a scavengedcarcass-were being changein the hominid line leading to reservedfor group consumptionwhen full humanity over the past two million the foragingpartiesreconvened at their years has been the great expansionof startingpoint. languageand communicationabilities, To the observerin the time machine togetherwith the cognitive and cultur- behavior of this kind, takeDin context al capabilitiesintegrally relatedto lan- with the early hominids' practice of guage.What is the evidencein support making tools and equipment, would of this surmise? seemfamiliarly "human." If, as I supOne humble indicator of expanding pose, the hominids under obseryation mentalcapacities is theseries of changes communicated only as chimpanzees do thatappears in themostdurable materi- or perhaps oivery rudimentaby means al recordavailableto us:the $tonetools. !y protolinguisticsiglals, then the obThe earlier tools from the period under servermight feel he was witnessing the consideration hereseemto me to showa activitiesoi somekind of fascinating bisimpleand opportunisticrangeof forms pedal ape. When one is relying on arthat reflect no more than an uncompli- chaeology to reconstructprotohuman catedempirical graspof one skill: how life, onemust stronglyresistthe temptato fracturestoneby percussion in sucha tion to protect too much of ourselves way as to obtain fragmentswith sharp into the past. As Jane B. Lancasterof edges.At that stageoi toolmaking the the Universityof Oklahomahaspointed maker imposeda minimum of cultural- out, the hominid life systems of two milly dictatedforms on his artifacts.Stone lion yeals ago have no living counterperform perfect- parts. toolsassimpleasthese ly well the basicfunctionsthat support progressin the direction of becoming SocialAdvances human, for example the shapingof a diggingstick,a spearand a bark tray, or My model of early hominid adaptathe butcheringof an animal carcass. tion can do more than indicatethat the The fact is that exactly such simple first toolmakerswere culturally protostone tools have been made and used human. It can also help to explain the ever since their first invention, right dynamics of certainsignificant advances down to the presentday. Archaeology in the long course of mankind'sdevelopalso shows,however.that over the past ment. For example, one can imagine several hundred thousand years some that a hominid social organization in-

Clark of the University of Califomia at Berkeleyand Lewis R. Binford of the Universityof New Mexico.In their view the earliestmeat-eaters might have obtained the fleshof animalsweighingup to 30 kilograms by deliberatehunting, but the fleshof larger animalswasprobably availableonly tfuough scavenging.



ing implicationis that in eachsuccessive generation the more capableindividuals made better tools and thereby gained advantages that favored the transmission of their genesthrough natural selection: it is supposed that thesegreater capabilitieswould later be applied in aspects of life other than technology. Another hypothesis regardshunting as being the driving force. Here the argument is that hunting requiresintelligeoce,cuDning,skilled neuromuscular coordinationand. in the caseof group hunting, cooperation. AmongothersugAccounting for Evolution gested prime moversare suchpractices Thus the food-sharing hypothesis as carrying and gathering. now joins other hypotheses that have If we compare the food-sharingexbeen put iorward to account for the planation with thesealternativeexplacourse of human evolution. Each ol nationswe seethat in fact food-sharing thesehypotheses tendsto maintain that incorporatesmany aspectsof each of one or another innovation in protohu- the others.It will alsobe seen that in the man behavior was the critical driving food-sharing model the isolated eleforce of change. For example,the argu- mentsare treatedas beilg integralparts ment hasbeenadvanced that tools were of a complex,flexiblesystem. The modthe "prime movers." Here the underly- el itself is probably an oversimplified

pasttwo million yearsor so. I have also set forth alchaeologicalevidencethat food-sharing was an establishedkind of behavioramongearly protohumans. The nolion is far from novel:it is implic" it in many philosophical speculations and in many writings on paleoanthropology.What is novel is that I have undertaken to makethe hypothesis explicit so that it can be testedand revised.

versionof what actually happened, but il seems sumciently realistic to be worthy of testingthrough further archaeological and paleontological research. Lastly,th food-sharing model can be seento have interconnections with the physical implications of fossil hom!nid anatomy. For example, a prerequisiteoffood-sharingis the ability to carry This abilily in turn is greallyfathings. cilitated by a habitual twolegged posture. As Cordon W. Hewesof the University of Coloradohaspointed out. an important part of the initial evolutionary divergence of hominids from their primate relatives may have been the propensity andthe ability to caffy things about.To me it seems equally plausible pressures that the physicalselection that promoted an increase in the size of the protohuman brain, the!by surely enhancingthe hominid capacityfor communication. are a consequence of the shift from individual foraging to food$haringsometwo million yearsago.




With on Introduction by

Glynn Isaac
Univercily ol Colifonio, Eerkeley

Richard E. F. Leakey
Notionol Museumsof Kenyo

W. H. Freemanand Company
Son Froncisco

All of tle ScrrNrrrrc AMERTcAN aticles in llrman Ancesto6 aie available as separateOffprints. For a complete list of articles now available as Ofiprints, write to W. H. Freemanand Company,660 Market Street,San Francisco, California,94104. Library of Congress Catalogingin PublicationData Main entry imder title: Human ancestors. Bibliogaphy: p. Includesindex. I. Human evolution-Addresses, lectures. essays, 2. Paleolithicperiod-Addresses, essays, lectures. I. Isaac,Glynn L., 1937- II. Leakey,Richard E, III. ScientiffcAmerican.

cN281.H847 573.2 79-4486 ISBN 0-7167-1100-1 ISBN 0-7167-1I01-x pbk. Copyright O 1960, 1962, 1964, 1966, 1967, 1969, 1974, 1977,1978,1979by ScientiffcAme can, Inc. No part of this book may be reproduced by any mechanical, photographic, or electronicprocess, or in the form of a phonographic recording, nor may it be stored in a retrieval system, transmitted,or otherwisecopiedfor public or private use,without wdtten permission ftom the publisher, Printed in the United Statesof America 98765432