Evolutionary Anthropology 153

ISSUES

Diet and Food Preparation: Rethinking Early Hominid Behavior

SONIA RAGIR
At least four innovations in nutrient sources, preparation, and distribution underlie the transformation of our evolutionary ancestors into modern humans: the digging, preparation, and consumption of tubers and rhizomes; the technological mediation of hunting and butchering of animal prey; the socially mediated redistribution of animal prey; and the control of re for cooking. The emergence of technologies for processing hard-to-obtain or difcult-to-digest foods such as animal protein, savanna tubers, rhizomes, and perennial bulbs probably precipitated changes in energy availability and expenditure that directly affected gut proportion and indirectly affected sexual dimorphism, fetal weight, and brain size.1,2 The indigestibility or toxicity of several potential food items, including rootstocks and carrion, could be extremely important for understanding hominid adaptations to the mosaic ecology of late Plio-Pleistocene South and East Africa.35 Primate consumption of rootstocks, seeds, and meat, which is often assumed to be essential for hominid adaptation to the savanna-woodland, is constrained by digestion-inhibiting agents in underground storage organs and by the rapid bacterial contamination of carrion. These potential restrictions on hominid food choice have led me to argue that hunting rather than scavenging, together with technologically assisted extractive foraging and food preparation, was an essential parts of the earliest hominid behavioral repertoire.4,6 The archeological and fossil record and studies of comparative gut morphology demonstrate that major technological innovations and relatively minor adaptations in digestive physiology often accompanied new sources of nutrition.1,7,8 Because plant toxins and digestion-inhibiting compounds constrain primates dietary choices,5 their presence in a majority of savanna rootstocks and mature leaves must have had an impact on food choices and motivated hominid food preparation.9,10 With the exception of wooden digging sticks, technologically assisted foraging is not usually attributed to Australopithecines. Yet some simple extractive techniques such as crushing, soaking, and termite shing can be found among chimpanzee and other primate communities, which suggests that, at the very least, these techniques would have been within the capacity of the earliest hominids.1113 Tubers, rhizomes, and perennial bulbs are rich in carbohydrates but low in protein, and may have been important sources of energy for early hominids.7,10 The edible wild tubers of the rainforest and savanna are mostly yams (Dioscorea spp.); when cooked, they become a staple food for extant populations of hunter-gatherers.10 Because their large size made feeding among the small ends of branches difcult, hominids frequently would have foraged on the ground where the vegetation from edible tubers and rhizomes grows out of the moist soil of a rainforest. Aboveground visual cues to the presence of edible roots would have been part of the knowledge that hominids carried from the tropical forest to riparian woods and out onto the baked ground of the grasslands.14 The pattern of wear on australopithecine incisor and molar surfaces and on the polished ends of long bone and antler tools at Swartkrans in South Africa has been interpreted as evidence of the extraction and consumption of rootstocks by Australopithecus and Paranthropus.15,16 The exploitation of mosaic, semiarid, and seasonal habitats by early hominids would have required either physiological or technological adaptations in order to include wild tubers and bulbs as staple items in their diets. Most apes cannot digest large quantities of toxic plant compounds. In contrast, herbivores have specialized chambers of the digestive tract in which a symbiotic intestinal ora deactivates plant allelochemicals through the process of fermentation. Chimpanzees and other large primates rarely exploit the underground storage organs that are the staple of many tropical hunter-gather populations. Outside the rainforest, in the savanna-woodlands of subtropical Africa, fewer varieties of tubers and bulbs are present.9 And most tuberous species in these drier areas have increased amounts of digestion-inhibiting compounds such as invertase, amylase, and proteinase inhibitors, which prevent the enzyme decomposition of fat, sugar, starch, and protein.3,5,17 Before the use of re for cooking, hominids would have had to remove or deactivate the digestion inhibitors in rhizomes and rootstocks mechanically by crushing, soaking, fermenting, or drying them, or by eating them with clay.4,6,10 Crushing and soaking leaches out allelo-chemicals in roots, while fermentation allows wild yeast to break down the remaining toxins. Pharmacological research has demonstrated how alkaloids and other toxic compounds in roots and rhizomes are effectively bound by clay. Humans deliberately use the adsorptive properties of clays and, occasionally, other substances such as charcoal and lime to bind toxins in food in ways that appear to be an elaboration of the geophagy that is observed in animals. The most common and reliable subsistence strategy in tropical and midlatitude modern hunter-gatherer pop-

154 Evolutionary Anthropology

ISSUES

ulations is based on a high daily consumption of roots, fruit, owers, and young leaves that are gathered primarily by women and children. These are supplemented with animal protein from both small, slow-moving animals, also collected by women and children, and larger mammals that are hunted by men. The proportion of animal protein in diets increases as hunters move to higher latitudes: meat constitutes 5% to 20% of the tropical hunter-gather diet but more than 70% of the winter diet of subArctic and Arctic hunters.18 As is true among most primates, there is a signicant difference in foraging range and diet between females and males in contemporary hunter-gatherers. It seems likely that this traditional asymmetry in the distribution of animal protein was also to be found among female Australopithecines and early Homo. Female chimpanzees tend to eat more low-nutrient foods than do adult males. Females also have to spend more time foraging than males do. Among chimpanzees, it has been observed that the use of tools for the exploitation of insects and hard-to-process oily nuts is primarily a female and juvenile activity.1113,16 Because rootstocks are hard to obtain and require time-consuming preparation in order for humans to absorb their nutrients, they most likely were gathered, prepared, and eaten by early hominid females and juveniles.18,19 It seems reasonable to assume that hominid adult males would have had relatively little need to exchange meat for plant food with females. Because of their larger size, they, like chimpanzee males, would have had preferential access to easily obtained and digested ripe or dried fruit, nuts, and meat during the wet and dry seasons. Because digging, as well as crushing and soaking, was necessary to obtain and detoxify rootstocks, female hominids may have played a central role in the emergence of tool use and manufacture.20 Until tools became necessary for processing foods in male-dominated domains such as hunting, australopithecine and early Homo food preparation and tool-making may have been largely female activities to gather hardto-obtain foods.12,13,20,21 Because of maternal provisioning during their relatively long period of semi-dependency, male juveniles

would have been familiar with tool use and could have transferred this knowledge to male activities such as skinning and butchering. However, adult males probably did not enter into the systematic manufacture of tools until early Homo erectus began killing and butchering larger herbivores.1,20 Killing large animals required implements for holding, battering, and piercing, while cutting and chopping implements were needed for skinning, deeshing, and dismembering. Males might have adapted the female tool kit. For example, they may have used sharpened digging sticks to pierce and hold prey, pounding and crushing stones to dismember carcasses and extract marrow, and root peelers and scrapers to skin and de-

Because digging, as well as crushing and soaking, was necessary to obtain and detoxify rootstocks, female hominids may have played a central role in the emergence of tool use and manufacture.

esh carcasses. Among ethnographic hunters, hunting and afliative exchanges are time-consuming and leave the men with little time or motivation for other foraging activity.21 As early Homo erectus males engaged in hunting larger prey and extended sharing of meat, they too would have had less motivation for foraging or preparing plant foods. The larger herbivores found butchered in East African Acheulian sites suggests that more meat was available than the hunters could consume in one sitting.1,8,13 An institutionalized exchange of meat and prepared plant food between males and females may have commenced and intensied among Homo erectus. Demand sharing rather than formal reciprocal exchange could have provided the foundation

for alliances among the males. These male alliances and female provisioning of semi-dependent juveniles could have created strong enough extendedfamily and community ties to support closer birth intervals and larger groups. The existence of this yearround supply of protein that is essential for the growth of a large brain and body not only supported changes in fetal and postnatal development, but also fueled population expansion and the dispersal of H. erectus out of Africa. On the other hand, the long-term reciprocal exchanges that underlie regional intergroup networks could hardly have emerged on the basis of demand-sharing of an episodically available and rapidly spoiling resource like meat.21 Intergroup alliances such as marriage, regional-networks for exchanging raw material and nished products, and social stratication, which depend on longterm reciprocity, would require food preservation, storage technologies, and a more elaborate division of laborthat is, a specialization in the distribution of knowledge and production skills.2 The transition from late presapiens to archaic H. sapiens some 300,000 to 250,000 years ago was accompanied by the abundant evidence of the use of re for cooking.22 A revolution in food preparation, storage technology, and food distribution was precipitated by the use of re. Because hominid females probably were the main participants in technologically assisted food extraction and preparation, and rarely strayed far from the safety of the social group, they were also available to maintain res once a home base had been established. The sexual differentiation of gathering and hunting activity and the tendency of nonhuman and human females to forage close to the main group or camp put hominid females in a position to control the use of re. Thus, it is likely that women were responsible for cooking and the innovations in human culture that followed from it.4,13 Cooking also gave females access to meat above and beyond those portions that they received after the hunters had shared with other males. In the process of cooking, females could informally appropriate pieces of meat for themselves and their infants before the formal redistribution of

ISSUES

Evolutionary Anthropology 155

choice foods, which probably was still dominated by hunters and male-male afliation. The enormous cranial volumes found in archaic Homo sapiens and the reduction of size dimorphism to modern proportions in late Homo erectus may be explained in part by an animal protein component in the female and juvenile diet comparable to that of modern high-latitude hunters.18 This last signicant episode of encephalization, the one that catapulted presapiens to Homo sapiens, may best be understood as the result of intensive hunting of large game and a cultural revolution that intensied with the use of re for cooking.1,4,22 Female hominids were likely to have been the innovators of the early plant processing technology that males transferred to hunting. Hunting larger animals may have initiated demand sharing of meat and a greater year-round dietary stability. Nutritional stability in females then led to shorter birth spacing, and changes in the number of dependent juveniles in communities. This shift in the age composition of the group paved the way for group responsibility for the socialization of semi-dependent juveniles. The growth in population probably initiated hominid expansion into the vast temperate plains and forests of Europe and Asia. Concurrently, the increasing density of late Homo erectus and early Homo sapiens populations may have been the impetus for long-term reciprocal exchanges, insuring more balanced, hierarchical relationships within communities and stable relationships between them. Because of their inherent advantages over females in size, strength, and access to preferred foods, as well as their lower expenditures of reproductive energy, hominid males were likely to have been behaviorally conservative. They did not have the motivation to invent nor would they have readily adopted energy-expensive technological innovations. Why should they? Preferential access to the high-nutrient foods that were easy to obtain, process, and digest did not require technological mediation of male foraging or hunting except, perhaps, in times of severe ecological crisis or population growth.

A conservative interpretation of the evidence from primate behavior leads to the assumption that, as among chimpanzees and baboons, female Australopithecines rarely proted from male hunting activity. Australopithecine females, who were meeting the protein and energy requirements of encephalization in fetuses and semi-dependent juveniles, were not only capable of but also motivated to develop the technology for digging, crushing, and detoxifying rootstocks. The association of early hominid tools and early Pleistocene African kill sites suggests that hominid males clung to opportunistic hunting without elaborating its technology for more than two million years. Finally, about oneand-a-half to two million years ago, male hominids probably borrowed and adapted the female foraging technology to satisfy an increased demand for animal protein that arose as the brain enlarged, the birth interval decreased, and the proportion of semi-dependent juveniles increased. The requirement of afliative exchange between adult males and an increase in the number of semi-dependent juveniles probably motivated males to practice more intensive cooperative hunting of large prey, and permitted females access to a relatively stable supply of protein and amino acids essential for reproduction and growth.

ACKNOWLEDGMENTS
I thank Nancy Bogen, Department of English, Sandra Brandler, Department of Sociology, Anthropology, Social Work, and Martin Rosenberg, New York University Medical Center, for their criticism and comments.

REFERENCES
1 Aiello LC, Wheeler P. 1995. The expensive-tissue hypothesis: the brain and the digestive system in human and primate evolution. Curr Anthropol 36:199 221. 2 Deacon TW. 1997. The symbolic species. The co-evolution of language and the brain. New York: W.W. Norton. 3 Buonocore V, Silano V. 1986. Biochemical, nutritional and toxicological aspects of alpha-amylase inhibitors from plant foods. In Friedman M, editor. Nutritional and toxicological signicance of enzyme inhibitors in foods. Advances in experimental medicine and biology vol. 199:483507.

4 Stahl AB. 1984. Hominid dietary selection before re. Curr Anthropol 25:151168. 5 Waterman PG. 1984. Food acquisition and processing as a function of plant chemistry. In Chivers DJ, Wood BA, Bilsborough A, editors. Food acquisition and processing in primates. New York: Plenum. p 177211. 6 Gordon KD. 1987. Evolutionary perspectives on human diet. In Johnson FE, editor. Nutritional anthropology. New York: Alan R. Liss. p 339. 7 Lee-Thorp JA, van der Merwe NJ, Brain CK. 1994. Diet of Australopithecus robustus at Swartkrans from stable carbon isotopic analysis. J Hum Evol 27:361372. 8 McHenry H. 1994. Behavioral ecological implications of early hominid body size. J Hum Evol 27:77 87. 9 Peters CR, OBrien EM. 1994. Potential hominid plant foods from woody species in semi-arid versus sub-humid sub-tropical Africa. In Chivers DJ, Langer P, editors. The digestive system in mammals. Food, form and function. Cambridge: Cambridge University. p 166 192. 10 Johns T. 1996. The origins of human diet and medicine: chemical ecology. Tucson: University of Arizona Press. 11 Boesch C. 1993. Aspects of transmission of tool-use in wild chimpanzees. In: Gibson KR, Ingold T, editors. Tools, language and cognition in human evolution. Cambridge: Cambridge University Press. p 171184. 12 McGrew WC. 1977. Evolutionary implications of sex differences in chimpanzee predation and tool use. In: Hamburg DA, McCown ER, editors. The great apes. Menlo Park: Benjamin/ Cummings. p 441 463. 13 Boesch C, Boesch H. 1984. Possible causes of sex differences in the use of natural hammers by wild Chimpanzees. J Hum Evol 13:415 440. 14 Hunt KD, Nishida T, Wrangham RW. (n.d.) Sex differences in chimpanzee positional behavior, activity budget and diet: relative contributions of rank, reproductive-demands and body size, and implications for tool-use. Am J Phys Anthropol 15 Schoeninger MF. 1996. Stable isotope studies in human evolution. Evol Anthropol 4:8398. 16 Kay RF. 1985. Dental evidence for the diet of Australopithecus. Ann Rev Anthropol 14:315341. 17 Prathibha S, Nambisan B, Leelamma S. 1995. Enzyme inhibitors in tuber crops and their thermal stability. Plant Foods Hum Nutr 48:247257. 18 Hayden B. 1981. Subsistence and ecological adaptations of modern hunter-gatherers. In: Harding RSO, Teleki G, editors. Omnivorous primates. New York: Columbia University Press. p 344 421. 19 Hawkes K, OConnell JF, Blurton Jones NG. 1997. Hadza womens time allocation, offspring provisioning, and the evolution of long postmenopausal life spans: with CA comment. Curr Anthropol 38:551578. 20 Zihlman A. 1997. The Palaeolithic glass ceiling: women in human evolution. In: Hager LD, editor. Women in human evolution. New York: Routledge. p 91113. 21 OConnell JF, Hawkes K, Blurton Jones NG. 1999. Grandmothering and the evolution of Homo erectus. J Human Evol 36:461 486. 22 James SR. 1989. Hominid use of re in the Lower and Middle Pleistocene: a review of the evidence. Curr Anthropol 30:126.
2000 Wiley-Liss, Inc.