Earty hominid hunting and scavenging: role of meat as an energy source John D. Speth Museum of Anthropology, University of Michigan, Ann Arbor, MI 48109, U.S.A. the Perspectives have shifted recently from the traditional view that early hominids were hunters to one which now sees them as opportunistic scavengers. However, both views share the common underlying nutritional assumption that meat inevitably provides a “high quality” food that will he incorporated into human diet more or less to the extent that animals are available and accessible. This paper argues that meat may actually have been a relatively marginal source of sustenance for early hominids, because physiological limits to total protein intake (plant and animal), scarcity offat in most African ungulates, comparatively high levels of protein in many plant foods, and the inability of early hominids to extract lipids from the cancellous tissue of bones, acted together to maintain their total meat intakes at modest levels, particularly during seasonal or inter-annual periods of resource stress. Received 31 May 1988 Revision received 20 January 1989 and accepted 25 February 1989 Keywords: early hominids, hunter-gatherers, hunting, scavenging, human diet, protein, energy. Journal of Human Evolution ( 1989), 18,329-343 Introduction The diet and subsistence strategies of early for well over a century, and from have of this debate. been human at the center evolution separation game of early as the driving 1953; Washburn Hill, saw increased hominids hominids. taphonomic techniques, the foundation Gorge, Binford, australopithecines, which Torralba; 1987b; Klein, and greater and success 1969; Schaller have begun see Bunn, 1987). The result 1986; of these may not have been the inveterate large-game hunters hominids, perhaps as recently as the Late Pleistocene, entirely other by opportunistically predators & Crader, occurred adults (Einford, 1981; Jones, scavenging 1984; Potts, of larger While abilities archaeological of the classic erected later of and sites that (e.g., Olduvai & Stone, is a growing consensus pre-modern of large mammals 1986, 1987; Bunn 1982; Shipman, small 1982; 1986; that hominids, we once thought they were. Instead, may have obtained meat largely or the carcasses have been larger (Dart, 1969; Foley, 1982; Binford even 1984, 1987a; Blumenschine, at all, the prey would many new analyses perhaps of to the in taking the hunting new had been Potts, models leading 1987). re-examined Homo erectus, and perhaps efficiency to question hypothesis factor of all later hominids of sophisticated have the hunting and meat-eating accepted & Lowther, see Gordon, an arsenal anthropologists upon Zhoukoudien, primates, of ideas, the focus of lively debate as the principal the emergence anthropologists with been the most widely on hunting other review Armed have the role of hunting recently, 1968; Pfeiffer, general In the last few years, were from & Lancaster, pre-modern Until reliance force underlying 1982; for a recent hominids the very beginning 1983; Trinkaus, animals, and perhaps abandoned by et al., 1980; Isaac 1987). If hunting the young or sickly mammals. this shift in perspective marks an important step forward the early hominid archaeological record, both models-scavenging the common underlying assumption that meat inevitably constitutes in our ability to unravel and hunting-share a “high quality” food that will be incorporated into human diet more or less to the extent that animals are available and accessible (cf. Lee & DeVore, 1968; Winterhalder & Smith, 1982; Tooby & DeVore, 1978: 211; but see Hill, 1988). In brief, in this paper I attempt to show that meat 0047-2484/89/000329 + 15$03.00/O 0 1989 Academic Press l~imitcd 330 ,I. D. SPE’Il1 may at times in fact constitute there is a ceiling a relatively to the amount consume for calories on a sustained total calories under normal, during times ofresource the animals themselves or difficult to obtain, may be forced foragers, for their overall budgets, health, Thus, evolutionary models semiarid do during savanna habitats adequate non-protein few concrete important contribution others ofits importance; concerned daily and seasonal seasonal or inter-annual I can make here is simply efforts and creativity 1983, literature 1987 a, 1987b, on foragers, in press) seek out fatty meat and marrow fat is a highly concentrated especially research metabolized ofscholars protein intakes kcal per gram, while protein and carbohydrate There have vet-v will find ways to address it. and ethnohistoric to selectively other foods do not. Moreover, 1986; Speth, of access to to raise the issue and try to convinct 1 am confident that the collective 1983; Speth, efficiently hominids for how we should proceed to address this issue. Perhaps the most of hunters vitamins energy stress in the 1 confess at the outset that I presently 1981). Fatty foods not only taste good but they produce more of how early periods of resource tendency fat-soluble consequences work patterns, Africa to assure themselves Hayden, also times present are missing a critical and central issue: what of eastern and southern the ethnographic & Spielmann, Such patterns. Maximum (Speth v~hen total caloric intakes, protein. that focus only on the question with human evolution In exploring adequate of dietary Africa), and edible plant mods scarce (Lt’iens, 1977), with important body weight, sources of energy? suggestions sources, either fat or carbohvdrate. to maintain amounts meat (i.e., by hunting or scavenging) did early hominids specilic;lll~. means that halfor more ol‘a forager’s lean or fat-depleted “ bottlenecks” and fertility and mortality obtained conditions, in their attempt stature, More that a forager can sali.ly- stress (i.e., the dry season in sub-Saharan excessive foragers with critical adaptive and plant) from non-protein may be extremely to consume (animal basis. ‘I’his upper limit, ahout 300 g or roughly XY% 01 non-stressful total energy needs must he obtained However, poor source of sustenance. of protein than and I (Jochim, 1981: 78-87; a feeling of satiety that source of energy, supplying 9 each provide only 4 kcal per gram. Fat is protein. and are sources of essential Spielmann noted the widespread Finally, fatty foods carry fatty acids (Guthrie, important 1975; Mead et nl.. 1987a). is another during important periods on the subject, nutritional of resource reason, however, why fat is critical to foragers, Although there has been relatively little stress. there appears to be an upper limit to the total amount of protein (plant and animal combined) that an individual can safely consume on a sustained basis without serious health consequences. This limit-most appropriately expressed as the total number of grams of protein per unit of lean body mass that the body can safely handle-is about 300 g or roughly 50% ofone’s normal total daily caloric intake (George F. Cahill,_Jr., pers. comm.). Prolonged protein intakes above this threshold may exceed the rate at which the liver can metabolize amino acids, and the body can synthesize and excrete urea (Buchanan, 1988; Noli & Avery, 1988; Speth, in press), leading ultimately to hypertrophy and functional overload of the liver and kidneys, elevated, or even toxic levels of ammonia in the blood, dehydration and electrolyte imbalances, severe calcium loss, micronutrient EARLY deficiencies, and lean tissue HOMINID HI’NTING loss (Cahill, 3’I 1 AND S(:A\-ENGING 1986: 42; Hegsted & Linkswiler, 1981; McArdle r/ 545; Miller & Mitchell, 1982: 115-l 16; Whitney & Hamilton, 1984: 145). remain poorly documented and The precise nature and value of this limit, however, controversial. Laboratory rats, often used as analogues in studies of human diet and nutrition (see discussion in Hill, 1988), are able to adapt to diets that provide greater than al.. 1986: 50% of the energy time required makes as protein, is much the interesting although longer at protein (Alfred observation levels above E. Harper. pers. about comm.). 35%, the adaptation Harper (pers. 1 comm. that: \vhen rats are lirst offered a high protein diet their li)od intake decreases, their ,growtJl decline\. increases very substantially but, after ;1 period of adaptation, thr. rntt’ :tt water consumption which they can metabolize amino acids from protein increases greatly. As this occurs. their ii~~l intake increases, their growth rate increases, and the)- seem to perform almost as WYII as those orqanz receiving a moderate amount of protrin (Anderson rt d., 1968). I,iver and kidney. intimately involved in amino acid metabolism and elimination of waste products liom protein, hypertrophy (Munro. 1964). The adapted animals sure-ive \veII, although there is somr evidrncc that their longevity may be shortened. Interestingly, if rats that are well adapted to extremt$ high protein diets are offered an opportunity to choose betbveen low and high protein diets. thr) will select from the two to obtain about 23-35% of their energy as protein and, over a short of encrxy as protein (Peters et(il.. 1983i period of time, this usually declines to between X-2.5% Similarly. dietary Castonguay ( 1981) obsetxrrd increasing which growth protein protein .Ilthough protein below (only protein similar levels IO days) on the order ;Inothcr Ibcusing basclinc ol)tain intakes containing rats 25% Hurkhardt content although selected Donald protein of the diet rates up to l~\x~ls irl were hi,yhe~t to question to obser1.e as well as other u-hether & Margen, for establishing (cf. Hegsted, Callowa), ill effects in healthy deleterious Regrettably, 1978: the test subjects & hIargen male elTects, the duration adapted 1979 fi)r observations o\.er time to on adaptation ( 1968; see also Harper? volunteers in of his who consumed in .I 197411: a purifir*d provided up to 70’%1 of total calories. However, most studies with to a safe upper protein threshold tying between these two cJ\trc*tnc.x. 50% of total ener,qy et al., 1986: 545; McGilvery, the probable of contemporar!. order human (c.<q.. CIahill, 1986: 42: Eaton St Iiol~nc~r, 1983: 791). ofthis of magnitude populations 10% and 15% of their total energ)’ limit is to look ;lt tht in various requirements cited in Gaulin & Konner, 1977: 571, in a global including a number of cases from northern latitudes. 17%; ilt levels. their appropriateness calories. (19.58; I!~~!W0 (median in obscr\~cxl parts of tht. world, particularly on those which routineI>, consume large amounts of protein. for comparison, McGilvery (1983: 793) notes that most human populations between :I rf /I/. than et a/. (1982) growth at higher is sub.ject of total extreme, way to determine protein the protein in the blood, 25-30% (see Oddoye g or about 1985: 287; hlcl\rdle t;inall>., in humans adult few studies ofverv high protein intakes have bet-n conducted are somewhat ambiguous. For example, Cheremin (198.3) was too short no obvious of300 rats fed diets of calories. lean in the form of protein. and fell off rapidly of ammonia At the other genetically rats are interesting, provided intakes 158) observed 25% lo-15% relativelv the results protein study). that of calories the higher about and tolerances diet in which protein human subjects point actual in rats oflaboratory elevated in which these gain in adult about studies stlldy weight rates 62). Unfortunately, in humans, and 30% diet with only 5% protein. requirements observed over contributed levels found (1982) slightly greater rats fed an isocaloric diets et al. mix containing range 8-50%). d somewhat from protein. As ;I t&a\ \Vhitinq survey of the diets of I 18 socit.tic*s. arrived at an average figurtb 01 tii,qher \aluta than hlcGil\.er1,‘s. 332 J. D. SPETH Since the diets of many inadequate (e.g., Phipard, intakes ofjust generally Third W’orld populations the better-fed populations in industrial fall below about 15% of calories: American workers 12.5-12.8%; & Young, Japanese 14.4%; 11.1% a better idea of the upper limits of protein came into vogue, many athletes before strenuous competitive intakes, expressed already noted. Here too the percentages 12%; Italian shipyard (McGilvery, as percentages proportions of protein, ranging diet (Shils & Young, 1988: Games (Steel, in Mexico, 1970: 121). athletes can be obtained & Young, are not substantially between higher than those 17% and 18% of their 1988: 1012). Similar intakes were observed 1988: 1012). Russian athletes the average daily protein intake In an early study of Olympic was only athletes, Lemon et al. (1984: 482), summarizing conclude about 80% thus far exceed For example, of their per capita dietary and blood. Approximately -was Turkana recommended 1985: iv; Little et al., 1988: 724). During the dry season, stress, decline and the contribution Kenya 30% (Galvin, and comparable, northern averaging 1971). The Northern intakes 1985: 184-194). Tanzania Although Ache, foragers their of calories in the neotropical generally meat, by protein a period of of protein may available data for the Masai ofsouthern are less precise, less than 30-35% of Kenya energy from animal products-milk, (Galvin, on of total calories. in northern resource approach 12%-15% 2 1% of their total calories on average are provided their total calorie for the literature about 20% pastoralists of total et al. ( 1949) that a diet in which protein provides calories is adequate for most. None of the protein values summarized Higher values do occur, however. 14.4% Berry A somewhat higher value-20% (1964). intakes by athletes, took in lower from 11% to 13% depending on total calorie levels of the 1012). A mong Australian athletes competing in the 1968 Olympic by Jokl and values for their protein reported a mean intake of 16.6%. athletes by loading” of meat while in training the available consumed (Shils & Young, (Shils consumption large quantities Nevertheless, of total calories, Italian calories in the form of protein obtain 1983: 790: Shils In the recent past, before “ carbohydrate consumed events. For example, in a study of weight-lifters nutrient or the protein 1988: 1011). Perhaps calories nations. marginal to consider 14%; Swedish West German looking at the protein intakes ofathletes. Olympic today are considered 1974: 178), it may be more instructive protein (cf. Taylor intakes appear to be & Ho, 1971; Ho et al.. forests of eastern Paraguay, consume considerable quantities of meat (approximately 56% of calories), and their average daily protein intake (including plant protein) is about 39% of calories (Hill & Hurtado, in press; Hill, 1988: 178). By far the highest per capita daily protein intakes are found in the arctic among Eskimos and other circumpolar hunting populations. Unfortunately, there are relatively few detailed studies, and most deal with groups already at least partly acculturated to western diets. Nevertheless, the results help pinpoint the upper limit of protein intakes. arctic population come from a Perhaps the lowest values recorded for a “ traditional” nutritional study of Lapps in northeastern Finland. This study revealed protein intakes ranging from 15% to 17% of calories (Hasunen 257). In northern Norway, migratory & Pekkarinen, reindeer herders obtained 1975; cited in Draper, from 15% 1980: to 23% of their energy from protein (Gassaway, 1969; see also 0grim, 1970). In Alaskan Eskimo diets studied in the early 1970s protein contributed between 22% and 25% of calories (Draper, 1977: 311). This same author suggests that the proportion of EARLY protein in the Alaskan premodern Eskimos, the contribution slightly over 45%. Coastal area by Angmagssalik protein & Krogh intakes that seen in other extended that under protein (McClellan 1944b, can thrive explorer, and within This where between 1936 and 1937, had Finally, Sinclair 56, 133). (1855) to total study many Euro-American similar had been to flourish of calories, all-meat the with in New York diet. discomfort. intake, of protein was deliberately Stefansson experienced The protein his condition replicated of calories, content returned not have survived for ill effects demonstration City that in which for an entire Details year of his diet reveal the remaining 75% raised reduced to nearly diarrhea, to about in 45% of calories. loss of appetite, 25% of calories, to normal. Stefansson’s with capacity et al., 1931). At one point nausea, was again meat and genetic fat providing et al., 1930; McClellan 74) diet, as 45%. by an experiment on an all-meat 25% (1953: no apparent clearest 44%. Eskimo as high explorers diet is provided in the same on a diet ofjust to a unique diets Perhaps conducted of Greenland calories ability 45-50% about of studied striking 1930; McClellan in et al. (1976) 1972 by Bang lived supplied diets approaching 1940, 1956). Eskimo 24% to a maximum (1908) supervision he was well acquainted lean meat in 25% to intakes on an Eskimo-like the proportion experiment studied Stefansson, generally 2 days from about Vilhjalmur At this level of protein and general 1960s for Canadian ranged close medical & DuBois, the experiment, on from about 01 protein of protein arctic values study reported century However, in the 1944a, (1928-1929) ofprotein In an earlier study (Herygaard, approaching 32%. An earlier 14, 44) the Eskimo’s populations. (Stefansson, about a range Greenland a mid-19th intakes Euro-Americans the arctic 43% from periods higher, in the early from protein. 333 SCAVENGING on the east coast ofGreenland, to ascribe fat, with protein AND to total calories the contribution It is tempting was collected (1914: of nearly data concluding calories Eskimos re-evaluated diet in northwestern of their Krogh data of protein Eskimos 26% HIJNTING et al. (1962) o b served of 29%. presents which derived Eskimo Mann 38% with a mean Schaefer (1977) HOMINID (1944a: the nutritional 234) firsthand hazards experience of an imbalance in the Arctic, in the ratio of to fat: if you are transferred suddenly from a diet normal in fat to one consisting wholly of. [lean meat] you eat bigger and bigger meals for the first few days until at the end of about a week you are eating in pounds three or four times as much as you were at the beginning of the week. Hy that time you are showing both signs of starvation and protein poisoning. You eat numerous meals; you feel hungry at the end of each; you are in discomfort through distention of the stomach with much food and you begin to feel a vague restlessness. Diarrhoea will start in from a week to IO days and will not be relieved unless you secure fat. Death will result after several weeks. Stefansson merits (1944b, of pemmican, 1956) a mixture also wrote extensively of fat and powdered on the jerked composition (dried) bison and nutritional or caribou meat (occasionally, though by no means invariably, mixed with small quantities of fruit). Eighteenthand nineteenth-century Native Americans, buffalo hunters, explorers, and trappers throughout the North American plains and arctic relied for months at a time on fur-trading companies became pemmican with no adverse effects. In fact, competing engaged in bitter staple (Stefansson, conflict-the “pemmican 1944b). Citing numerous over control of this critical frontier war”early descriptions of pemmican, Stefansson 334 J. 1956: 3 1, 2 12-2 13) argued ( 19441~: 90, every six or seven northern latitude Spielmann, pounds 1983), that the ideal mixture of fresh wild ungulates D. SPE’I’H was about lean meat. Since contains about 2 1% protein the proportion of calories lean meat in pcmmican one pound of most and 2-&“/o fat provided ol‘ti~t liar tcmprratc (SW and S[Jr’th by protein & is about 30-35%. Iluring World military utility outside pemmican of the merits of the arctic, for 9 weeks. provided roughly of the calories. he switched “unease” and the medical below of this brief survey studies intakes noted 50% during oscillate fluctuations intakes protein 1974a, where intake elevated any all-meat diets 50%. or East to maintain 1974b). threshold, forth water is scarce water intakes (or other “traditional” pastoralists) where their protein intakes below about protein Such such acid to arid safe level of intakes requirt 1 am not aware as those contribution rapid amino the upper diets shown among when semiarid high protein comm.). the concern 50% inhabiting of intake lower elevated pers. the protein protein gorging. to obtain, high-protein of the conclusions stress, and because E. Harper, a feeling is probably to sharply foragers 40% At times, to the lower-protein ofresource scarcity on protein about produced diets parallel or dificult (1940: 56), in fact, documents one back in [act, 50% ofcalories (Alfred entirely mixture. limit to sustained to adapt for of its provided protein this periods between Moreover, be well below African HDygaard actual or inter-annual and and to a safe upper time for the body drinking will again substantially foragers back insufficient (Harper, The protein he changed levels in human and point seasonal rapidly allow environments, of protein the As a test of 1943 subsisted the lower mixture, until to comincc ration. of lean to fat, one in which in which using persisted earlier, of calories. especially blends and a second high-protein that 1) attempted as an emer,Tencv in the summer no ill rfl’ects to the “discomfort” about foragers, official two different 20% of the calories, however, 1956: 290-29 94, of pemmican an army He used He experienced pemmican. The results ( 1944b: IYar 1 I, Stefansson establishment seen regularly by Greenland of in Ache exceeds Eskimos of calories: The Angmagssalik Eskimos seem to prefer a diet which nearly covers half the energ! Larger quantities of either are only eaten with requirements by fat and half by protein. reluctance. It might be thought that the reason is that the Eskimos, \vho Irrquently have IL more blubber at their disposal than lean meat, try to save the latter by eating so much fat as possible. But this is not the only factor which lies behind the relatively constant proportion remained between the two groups of food constituents, as even in time of’ plenty the proportion approximately constant (in the case ofextreme hunger, howe\-et-, the consumption offat usualI) predominates). Despite the uncertainty that stilt surrounds the issue of high protein intakes, in subsequent discussion I wilt use a value of about 300 g ( roughly 50% of total per capita daily caloric intake under normal, non-stressful conditions) as the approximate upper limit to the amount which affected remain detailed of protein this threshold that varies can be consumed among foraging safely on a sustained populations because basis. The extent of genetic factors, to or is by the absolute amounts or relative proportions offat and carbohydrate in the diet, unknown. Clearly, input from medical and nutritional specialists, as well as more long-term studies of protein intakes among foragers in different ecological settings and under different dietary regimes, are critically needed to clarify this issue. EARL.Y HOMINID HITNTING AND S(:A\‘ENGING :3:.ii Protein intakes in San diet It is important to emphasize well as from protein meat. Most contribution levels of protein quality and foods, however, that of plants, than this limit studies probably does meat, digestibility applies to protein of hunter-gatherer because and because to animal many most protein are improverished diet plant plant from plant to ignore are inferior & Young, as is clearly fbods as or downplay the foods do, in fact, have proteins (Scrimshaw in protein, derived tend shown 1976). low-cr in biological Not all plant by the diet ofthp Knlahari San (Bushmen). According to Lee (1979: 271), on average over 16% of the total per capita dail), calorit~ intake of the !Kung (CU. 2300 kcal) is provided by protein. As Ixe himself notes, this is an high level of protein, exceeding the average intake of Americans ( 1-l%, ). and to levels (ca. 1 l-18%) seen in athletes in the days before “carbohvdratc unexpectedly comparable loading” came into vogue (McGilvery, 1983: 790; Lemon ef al., 1984; Shils 8r k’ounyi 1988: 1011). Despite from the high overall protein prOtCin from in meat, mOngOng0 is more or less without consuming alOne lower biological XCOUntS and ( 197.5) argues calories-are that actually 2- to S-year-old (1978: 62) concurs, 10% San’s to mret children, provided pointing that in their first 6 months calories in the form ol‘protcin, daily protein noting that ch uhta; as than those Sil berbaucr, in othrr protein the But comparatively g, pro\riding plants Central high on average Konner ut cd.. n.d.; prott,in from staples protein, li)ragrrs Eaton levels (12dair. protein total nuts 1985). al~out energy :i’% intake f&d, tht, I!#?: Howc~ <‘I‘. requirements, 01‘ total t~\-c~n iti is adcquatc. milk, obviously about an idc~al li)od providing onl!. the Irvel fi)und in rite, and _ content by the who !liung as the \\‘ehmeyer are not.just minimum It is worth nuts (Ric-inoddr-on (CU. 2 I%: S~~cbth & meat found baobab their foods alone. in mongongo arr lack exceed on plant of fresh XW% ~), 1980: 5.57~; levels commonly margin, of protein mj]k; Kalahari, protein 01‘ in mongongo breast of protein consumed 198 1: 274; Tanaka, a\‘erage protein content tubers. fruits) Pxploited of’ animal non-arctic (28.8%) (Sderocuryc7 rcquircm(~nt rrcornm~nd;ltion we ignort. or “. by a generous high ‘l‘ht c{Umtit!’ and corn.” by weight Comparably 3 1~6%). marula nuts ‘rhk protein of life, is a “low-protein” the average 1983). their the San probably perhaps the proportion 34~I%,). and se\,eral such of wheat are correct, requirements, mrtanenii) is higher Spielmann, content and Hegsted daily conit’s Ibods. if li)r thr moment daily out that human of total the protein and minimum infants If Payne CalOrieS. 1976: FAO/\VHO/LINU, lower protein intakes-only for growing less than in plant FAO/\VHO’s of the protein about 56% Oftotal minimum requirement, considerably Hcxgsted for at least 6% ofcalorics protein & Young, sufficient growing from if we accept digestibility 1978; Scrimshaw diet, only about comes an adult whatsoever, as the baseline quality in !Kung two-thirds to meet any meat 10% of calories Payne nearly nUtS adequate about 120; Hegstcd, level of protein while in tsin I~wns nuts (:l&~\onicr well as 1)). other (Hazthinitr di,qitcila; San aroul~‘;, (Imp, 197:J. l!~iO: mongon,qo r/ ul.. 1969). restricted to thcsc fi>w staples. ‘I%(. of over 150 species of veCgetable foods (c.,q.> roots, btxns. JIU~~. by the !Kung, Hadza, and Australian Aborigines rscecds S g/l00 over et al., 12% of the calories 1988: 78-79). such as mongongo the eclectic (c.,q.. Lee, plant 1979) \Vhilc nuts kood 01‘ in these tsin beans inventory alniost foods thy biological certainI\ (Eaton & Konncr. 1985: value and di,gcstibilit\ are undoubtedl) characteristic compensates of the in?rit,r San for an\ arid anlirio of‘ to tjl;rt othw acitl 336 J. D. SPETH shortages or imbalances in these or other vegetable that protein is a limiting staples. In fact, it seems highly unlikely factor in the diet of the San or other extant foragers. Excess protein and seasonal resource stress Ifwe accept the 300 g or 50% threshold sustained total daily protein intake, suggested above as the approximate and for the sake of simplicity presence of small amounts of fat in meat, the maximum consume on a regular basis, with absolutely upper limit to here we ignore the amount of lean meat the San can no intake of plant protein, would be slightly under l-4 kg per capita per day, providing about 1150 kcal or 288 g of protein. Ifwe include the protein from their plant foods, the San can take in, at most, only 40% of their calories (920 kcal) in the form of animal consuming protein, or about 230 g. This is roughly about 1.1 kg oflean meat. Because ofthe substantially must accompany varies markedly equivalent to greater water intake that a diet high in protein (animal or plant), and because San hunting success from day to day, their meat consumption be considerably lower. In fact, the actual averages onIy about 0.23 kg (Lee, per capita under normal conditions daily meat intake should of the !Kung 1979: 271). Given these figures, it is interesting to note an observation made recently by Lee: In late-spring and early-summer [late-dry-season and early-rainy-season] hunting camps the proportion of meat may sharply rise. In one study, the four hunters of a small camp of 12 killed 29 animals in 17 days for a per capita consumption of almost two kilos of meat per day. These bursts of meat-eating tend to be of short duration, however . (1982: 40. emphasis in original). This astounding meat intake would provide nearly 420 g of protein than 75% of an individual’s suggested total daily calories), earlier, and well above deleterious if sustained and elevated daytime anything (amounting to more a level well in excess of the 300 g limit recorded in the arctic, and therefore very likely for more than a brief period. As noted earlier, shortages of water temperatures at this time of year in the Kalahari are likely to exacerbate the dangers of gorging on meat, since water is not readily accessible away from the dry-season camps and the physiological need for a substantial increase in water intake does not elicit an immediate response (Alfred E. Harper, pers. comm.). Moreover, the apparent rapidity of the dietary shift from low to high protein intakes may exceed the rate at which the body can adapt to the increased amino acid load, again possibly leading to acute adverse effects (Harper, arising from heavy reliance 1974a: 16). Deficiencies on meat may compound in micronutrients these problems (e.g., vitamin Bs) (Alfred E. Harper, pers. comm.). The San apparently make considerable bones of their kills, carefully before discarding the carcasses returns, removing them (Yellen, appears efforts to extract the marrow 1977). Despite to be inadequate the lipids from the meat and and boiling the smashed-up bones these efforts, the total amount of lipids in to compensate for diminishing overall caloric and their body weights fall to their lowest values at the same time that their meat intake reaches its peak (Wilmsen, dry-season caloric intakes, 1978, 1982). Thus, in their attempt to maintain the San are forced to consume adequate an excess of dietary protein, a situation which may pose serious nutritional problems for them at this time of year, and especially during particularly bad years; reliable access, therefore, to non-protein sources ofenergy (i.e., animal fat, plant oils or carbohydrates) becomes a critically important issue. EARLY HOMINID HUNTING 337 AND SCAVENGING Protein, calories, and early African hominids The preceding discussion in early hominid foragers, hominid First, past or present, and origins or evolution. probably adequately micronutrient diet vitamin (e.g., been alleviated particularly years, with if large sources of calories, concerning economies stress. 1% or 2%, with deposits and ventral an average limb epiphyses cancellous otherwise viewed large-mammal periods of resource or scavenged, fat probably calories would would physiological ,4s I have ofenergy carcass fat content and northern remaining primarily critical as a source of calories takes of calories on a somewhat and current archaeological have been a prime also have been derived target, from and nutritional constraints already discussed, the total & Spielmann, tissue ofthe than data given the grease protein, would needed to of these the animals the latter appear its high caloric density. but here a number would have come into play. amount of protein that an early an inter-annual make the protein, in the not have the value or during look. Whether and processing the technology the dry season, different of the distal vertebrae for foragers 1977: 291-294), rather during 1968; ofstressed may drop below in the marrow lacked or condition, (Ledger, (Speth et al., 1976: 399). However, sources hominids 4% as issue seasonal the range of these animals the cancellous who as far as we know to early within in the brain, and within during even in prime latitudes fat levels in many primarily an interesting of less than This falls squarely on the diet in any significant ungulates, hominids, stress, hominids and maniac. were exploited as sources African et al., 1988: 443; Yellen, primarily hominid ungulates (see Bunn resources hunted early they probably raises 1983, 1987a; Crawford to early have It seems dependent foods like cassava however, 1987a). border, one of the most lean animal been available boil the bones. When offat (Speth, tissue, easily by early conclusion, in temperate mandibular foods. on a vegetable small prey. mammals for factor in ancestors 1978) could overwhelmingly plant into Most 1983). And by the end of the dry season, limbs of large low-protein of most African ungulates nutrient plant reliance easily captured or scavenging This 1985: 285; Speth, starving the value of meat by consuming heavy & Busse, or other incorporated protein. with just from 1976; Hamilton or by hunting, of resource lean, & Konner, needs arisen lizards, of starchy, were not concerning protein or micronutrient shortages. In fact, protein may limiting nutrient in human diet in the past few thousand by scavenging periods protein have of agricultural range the suitability are notoriously basic might the hunting mammals whether inter-annual to have been a limiting insects, the emergence quantity, or even that of a narrow Thus, issues is unlikely Bi2; see Hausfater, by consuming a response to either become an important cultivation interesting their shortages unlikely represents only have several protein probably played little, if any, role as a selective Under most circumstances, our early hominid fulfilled Whatever Eaton has raised adaptations. more were likely. Obviously, of important hominid could consume on a sustained basis as a source of calories, as in modern humans, was probably limited to less than about 50% of daily energy needs (it is possible, in fact, that the limit was substantially lower adapted smaller to much Moreover, drinking water during in small-bodied periods was limited, overall early hominids quantities of food shortage and if meat intakes the course of the period of stress, below a third of total calories. the actual whose ancestral diet was probably of meat). and declining fluctuated ceiling would total caloric frequently have been intakes, if access and significantly even lower, to over perhaps 338 J. 0. In addition, component would il‘ protein-rich of early suggest, have further consumed their protein higher the the diet traditional and metabolic words, rises for every compensate entirely diets, “specific both is less than that substantial, could that of either or animal) very li,rager\ and Gould have been safcl!. high are accompanied the greater Stefansson’s in protein, were by metabolic of 20%; calories ingestion its experimental accompanied in excess additional By comparison, fat or carbohydrate, in the diet, or SDA values) 20 or more in metabolism. of fat or carbohydrate w(trc‘ an iniport;lnt and modern been 16). For example, action” 100 kcal of food ingested, for the increase have protein (plant 1987a: dynamic nuts primates et al., 1984). of protein (Speth, and higher of animal of protein Eskimo (i.e., as seeds of both may at times amount efficiency cost to the forager such 1981; Peters proportion metabolic lhods as studies contribution & O’Brien, the metabolic the diet, constrained (Peters Finally, and plant hominid SP’E’I’H were needed of foods increases by in other to composed (SDA values) of only 4-6%. Thus, during times of declining total energy intake, the metabolic costs of fulfilling those needs primarily with lean meat become greater the higher the total meat component Lee’s of the diet. (1982) per capita description daily the view being developed foragers a poor with (cited consumption consequences. It should Recent Tanzania for them work approached be remembered, may, intakes of sustenance, lowest body one weight et al. (1988) by O’Connell at that time ofyear would provide deleterious health according to Wilmsen (1978, at this time of year, despite the high meat and Bunn et al. (1988) the position where to contradict potentially that levels of the dry season, 2 kg, may appear with however, in fact, be stressful may also seem to contradict (1988: 452) note of San diet in the height here that high protein source 1982), the San are at their conditions above) of meat at times presented here. suggesting intakes. among that the Hadza For example, in Bunn et al. that the Hadza enjoy their peak period of hunting and scavenging large animals during the late dry season. At this time, they eat a lot of marrow but also prodigious quantities of lean meat. While body weight during than data the dry season among However, season the !Kung in both are heavily hominids. Bunn chopping and have not yet been published suggest cases, that their seasonal (Henry T. Bunn, Hadza processed pers. and !Kung, for their content (1977: 291) emphasizes placed in cooking these parts [axial the fact that pots and boiled the !Kung to extract James F. O’Connell, pers. or scavenged during a technology not available to “. . the routine allude of adults less pronounced hunted ofgrease, carcass photographs loss may be much comm.; carcasses et al. (1988: 443), for example, boiling for the Hadza, weight units] for grease. also smash . up bones Hadza ). the dry to earl) practice .” Similarly, so that comm. of Yellen the)- can be the grease. The boiling process is straightforward: chunks of meat and bone are placed in the pot with The meat will then be boiled for over an hour until well done and raw marrow ma) water. then be added to enrich the gravy. The meat is then eaten, the bones tossed aside, and the grav) drunk directly Thus, both !Kung from the cooking vessel. and Hadza place great emphasis on boiling grease from limb bones and vertebrae. Despite this critical source of fat, at least in the case of the !Kung there nevertheless are clear signs of food stress in the dry season, most clearly reflected in their declining body weights, but very likely also reflected, at least in part, in their low fertilit) and high rates of infant mortality (Speth, in press: Howell, 1986). \Yhat would happen the dr)- season small amounts to their eastern but of lipids overall questions to the “prodigious” if they were unable given perhaps suspect Africa would 1 in no way to imply technology, confronting of more reliable of year, what I suggest that less in happen to these environments during what would answers in the semiarid than and Hadza In fact, given thr are no clearcut mammals-certainly be unable but probably or during to make a more 01’ of stress. periods we observe I toda).; During early hominids such periods, would to carbohydrate-rich one. This bad years, with critical strategies adaptive be strongly vegetable in the USC of any meat. marginal particularly stressed. foraging access at all times obviously hominid USC of animals. or early hominids, foragers, that would of calories. alternative by boiling? be substantially that were themselves in which at hand, the use of large the dry season, intc,nsified, there of the !Kung from bones also be true of early mean a source true during consumption grease ungulates While year-round-would grease-rendering animals on meat? the information the same provide in most African reliance and southern meat to extract 01’ would stilt be especialI>, were obtained from selection pressure would ha\-t, “bottlenecks” (LViens, 1977) favored foods, would if meat absence Meat that would perhaps assure including them significant use of below-ground forms such as roots and tubers (e.g., Mann, 1981: 35; i’inccnt, 1985). ‘I‘hus, whether early hominids hunted or scavenged large mammals, I suspect they did so rclatiL4y infrequently and been not I see no reason lipids, predation them protein. or scavenging toward greater abIt. to estract opportunisticalll-. by Plio/Pleistocenc and greater the precious and hominids reliance on large from the cancallous lipids their to anticipate principal target an upwardly in sub-Saharan mammals, tissue would havc~ spiralling Africa at least ratt‘ 01’ that not until pushc(I thr\ wcr(’ “Hunting ancl of bones. Acknowledgements An earlier version Scavenging in Association people of this paper Early was presented Human for the Advancement have contributed No one other than of Science, at various myself, however, stages in the symposium at Evolution” the annual Boston, entitled meeting Massachusetts, to the development can bt. held accountable of the Feb. American 14, 1988. &Man) of the ideas presented for the precarious here. limb on IO which 1 have voluntarily climbed with saw in hand. I would like to thank Dieter Noli and Graham Avery who provided me with their paper “Protein Poisoning and <:oastal Subsistence” work [Jniversity Alfi-rd prior to its publication to my attention. of Michigan E. Harper arguments and in the paper more in line with current Bar-Yosef. Robert Serge in 1988; their Shishkoff translated George of the Cheremin F. Cahill, and ofTered many thinking J. 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