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Copyright University of Chicago, acting through its Press Dec 1999

Meat Eating and Hominid Evolution1

The role of hunting and scavenging in human evolution has been a subject of intense
anthropological interest for decades and has produced some of the most widely discussed and
debated research findings and interpretations in the history of the discipline. In spite of the
central place assigned to studies of meat eating, scholars from the different subfields of
biological anthropology rarely find themselves in the same room at traditional conference
venues, and therefore research data from one subfield rarely penetrate the others, To remedy
this and to discuss recent findings, a conference, "The Early Human Diet: The Role of Meat,"
was held October 2-5, 1998, on the campus of the University of Wisconsin, Madison. The
conference was sponsored by the Wenner-Gren Foundation for Anthropological Research
and was organized by Craig B. Stanford (University of Southern California) and Henry T.
Bunn (University of Wisconsin, Madison. The 18 participants were paleoanthropologists,
primatologists, archaeologists, and students of foragers and of carnivores, including some
researchers whose fieldwork and expertise overlapped two or more of these specialities'. They
had been chosen for their intersecting research on the role of meat in the early human diet and
had been asked a year before the meeting to prepare papers focusing on one of the following
topics related to this theme:

1. How is hominid behavior distinguished from natural processes in the fossil record? How is
hunting distinguished from scavenging in the fossil record?

2. What are the major costs and benefits of hunting for living animals versus scavenging for
carcass meat?

3. How do hunters hunt? What is the role of cooperation and communication during the hunt
for both human and nonhuman animal hunters?

4. What are the costs and benefits associated with meat eating compared with relying on an
herbivore's diet?

5. What is the nature of the variation cross-culturally in the nutritional importance of meat to
modern foraging people? When and why does meat represent more than just a source of
nutrition for modem foraging people?

6. What is unique about the pattern of meat eating in modem people compared with great
apes?

7. What is unique about the pattern of meat eating in modem people compared with early
humans?

8. What was the role of meat eating in the geographic radiation of the genus Homo?

9. What aspects of meat eating and foraging for meat may have influenced the evolution of
human intelligence?

10. What is the nature of meat sharing among modern primates and human foragers, and what
are the implications for early human sharing patterns?

The role that meat played in hominid origins was the central topic. How much was eaten at
different evolutionary stages, how habitat affected meat eating, how meat was procured and
how it was distributed, and how its archaeological signatures in the fossil record are recognized
were all topics of discussion. Although a revisiting of the Man the Hunter paradigm of the 19
6os (Washburn and Lancaster 1968) was not a goal of the conference, much discussion
inevitably focused on the importance of meat in human evolution, both through hunting and
scavenging and through meat-sharing behavior. it was clear that modem reframings of both the
Man the Hunter model and Glynn Isaac's food-sharing model (Isaac 1978) continue to be
highly influential in the subfields of biological anthropology that address meat eating in hominid
evolution.

PALEOANTHROPOLOGICAL APPROACHES

The only direct evidence of the behavior of our Plio/ Pleistocene ancestors is their fossilized
remains. Paleoanthropological approaches to meat eating in human ancestry have historically
involved the study of the bones of hominids, which tell us very little about carnivory, and the
bone assemblages of associated potential prey species, which may tell us much.
Archaeological approaches use stone tool evidence to build an empirically based picture of
early hominid behavioral ecology which, together with data on paleoenvironment, can provide
a perspective on the place of both living prey and scavenged carcasses in the early diet.

The role of scavenging versus hunting, a long-standing debate among human-evolutionary
scholars, was not a topic of intense discussion at the conference. Participants agreed that the
traditional either-or dichotomy between these two foraging modes is not a valid one and that
paleoanthropologists have moved on to studying the degree of scavenging or hunting on a
species-by-species or population-by-population basis. Stiner, for instance, argued that
Paleolithic Homo sapiens in Eurasia were both hunters and scavengers and that unequivocal
evidence for hunting in Europe first appears with archaic H. sapiens 250,000 years ago. Speth
presented information on Neandertal settlement and meat-eating patterns at Kebara Cave,
Israel, and questioned whether Neandertal use of space and hunting behavior differed
qualitatively from those of anatomically modern H. sapiens. Tappen addressed the monolithic
use of a Serengeti model for the evolutionary ecology of early Homo in the Pliocene; her data
from other, wetter sites indicate that scavenging has been overemphasized in reconstructions of
early hominid foraging ecology. Pickering presented a discussion of taphonomic issues
affecting early hominid meat-eating evidence and reexamined the Swartkrans hominid material.
He argued that large carnivores accounted for hominid representation at Swartkrans through
regurgitation and defecation rather than by active transport. Ancillary experimental studies of
captive carnivore consumption of primates were also discussed. Rick described a Holocene
high-altitude human hunting population that harvested vicufias. The local abundance of these
animals created an unusual situation of a lowrisk hunting strategy while providing
archaeological signatures useful for comparison with other, less abundant assemblages.

These diverse case studies were placed in perspective by the remaining papers that had a
paleoanthropological focus. Sept placed meat eating in dietary perspective by discussing the
other, more frequently consumed items in the early hominid diet, plants. While our ability to
model the entire dietary paleolandscape is still quite limited, Sept creatively estimated how a
Pliocene riparian forest habitat might have fed an australopithecine relative to a member of
early Homo. Foley showed that there may have been a major evolutionary/ecological shift that
corresponded to an increase of meat in the diet with the appearance of H. ergaster. Finally,
Walker presented a theoretical model for the recent increase in hominid brain size based on
dietary quality (a theme returned to by many participants) and linked the ability of genus
Homo, in particular H. erectus, to produce a large-bodied, large-brained neonate to
increasingly effective meat procurement by hunting.

LIVING NONHUMAN ANALOGUES

Although the fossils provide the only direct evidence for hominization, without living analogues
our ability to reconstruct hominid behavioral ecology would be severely limited. Such
reconstructions must be done cautiously and must bear in mind that modern apes are probably
quite different from their Pliocene ancestors just as we are from our own. But using primate
models provides a range of behavioral and ecological adaptations likely to have been present
in the last common ancestor before the pongid-hominid divergence. Van Valkenburgh used the
community ecology of living carnivores to investigate the ecological relationship between early
Homo and the large carnivores with which it was sympatric. She argued that since extant
carnivores prey heavily on other carnivore species in apparent interspecific competition, early
hominids would have faced a severe predation threat from their sympatric carnivores, which
would have viewed them as food competitors as well as prey. McGrew contributed data and
perspectives on the use of invertebrates by early hominids based on the harvesting and
consumption of a variety of insects by modern chimpanzees. He suggests that regular use of
this overlooked food source can rival mammal-meat consumption in nutritional importance.

Stanford and Rose presented comparisons of chimpanzee hunting and meat eating with those
of a related primate. Rose described her work with capuchin monkeys (Cebus albifrons),
which are voracious predators of small mammals and show some striking parallels with
chimpanzees and also major differences (lack of active sharing). Stanford compared the
meat-foraging ecology of chimpanzees with that of some modern foraging people and argued
that searching for meat as opposed to eating it opportunistically during plant food foraging was
likely to have begun only after the advent of efficient bipedal locomotion.

Milton and Schoeninger tackled the reconstruction of the early hominid diet on the basis of
those of modern higher primates. Milton argued that hominids overcame the constraints of
relatively inefficient digestive apparatuses among the hominoids by turning to meat in increasing
amounts. Schoeninger compared the diets of modern apes with that of some modern foragers
and argued that hominids were able to extract greater nutrition from their environments through
enhanced food processing, aided later by the use of tools.

FORAGER BEHAVIORAL ECOLOGY

Although the use of modern foraging people as analogues for earlier forms of humans has
come under much attack in recent years, the fact remains that tropical and subtropical people
living off the land with traditional technologies provide the best available windows onto the
dietary behavior of early hominids in similar habitats. The study of hunter-gatherers can also
inform us about meat-eating signatures in the archaeological record, as evidenced by Bunn's
paper on Hadza meat eating and the study of cutmarks. He showed that by I - 75 million years
ago early Homo was acquiring carcasses of ungulates by both hunting and power scavenging,
butchering the carcasses selectively and transporting them to favored central locations for
consumption. Hawkes also used the Hadza to understand the meaning of the carcass
ownership in order to infer patterns and purposes of the distribution of meat by foragers. She
brought chimpanzee meat sharing into the equation as well to contrast and compare patterns of
ownership and strategic use of the carcass by the two species. Alvard [who was unable to
attend) described a model for the origin of human cooperative hunting and reviewed the
literature on cooperation among foragers. Winterhalder presented a review of intragroup
resource transfer among social animals, pointing to the diversity of patterns observed and
arguing that complex forms of meat sharing likely characterized hominids at all stages of human
evolution.

A book from the conference with chapters contributed by the participants is currently in
preparation, to be published by Oxford University Press. The volume, tentatively titled
Meat-eating and Human Evolution, is being edited by Stanford and Bunn and is scheduled to
appear in 2000.

[Footnote]
1. (0 1999 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved
0011-3204/99/4005-0009$1.00.
2. The conference participants, their affiliations, and paper titles were as follows: Michael Alvard, State University of New
York at Buffalo, "Mutualistic Hunting"; Henry Bunn, University of Wisconsin, Madison, "The Role of Meat in Hadza Diet
and Society: Archaeological Signatures and Evolutionary Implications"; Robert Foley, University of Cambridge, "The
Evolutionary Consequences of Increased Carnivory in Hominids"; Kristen Hawkes, University of Utah, "Is Meat the Hunter's
Property? Ownership and Explanations of Hunting and Sharing"; William McGrew, Miami University, "The Other
Faunivory: Primate Insectivory and Early Human Diet"; Katharine Milton, University of California, Berkeley, "A Hypothesis
to Explain the Role of Meat-eating in Human Evolution: A Means of Painting One's Way Out of an Evolutionary Corner";
Travis Pickering, University of Wisconsin, Madison, and University of the Witwatersrand, "Taphonomy of the Swartkrans
Hominid Postcrania and Its Bearing on Issues of Fire Management, Meat-eating, and Extinction"; John Rick, Stanford
University, "Specialized Meat-eating in the Holocene: An Archaeological Case from the Frigid Tropics of High-Altitude
Peru"; Lisa Rose, Washington University, "Meat and the Early Human Diet: Insights from Neotropical Primate Studies";
Margaret Schoeninger, University of Wisconsin, Madison, "The Third Chimpanzee or the Fourth Ape: A Diet Perspective"
(coauthored by Henry Bunn, Shawn Murray, and Travis Pickering); Jeanne Sept, Indiana University, "Modeling the Edible
Landscape"; John Speth, University of Michigan, "Neanderthal Hunting and Meat-processing in the Near East: Evidence
from

[Footnote]
Kebara Cave (Israel" (coauthored by Eitan Tchernov); Craig Stanford, University of Southern California, "Hunting
Primates: A Comparison of Social Meat-foraging by Chimpanzees and Human Foragers"; Mary Stiner, University of
Arizona, "Carnivory, Coevolution, and the Geographic Radiation of the Genus Homo"; Martha Tappen, University of
Minnesota, "Rethinking the Behavioral Ecology of Plio-Pleistocene Hominid Meat Acquisition"; Blaire Van Valkenburgh,
University of California, Los Angeles, "The Dog-EatDog World of Carnivores: A Review of Past and Present Carnivore
Community Dynamics"; Alan Walker, Pennsylvania State University, "Neonate Body Size and Hominid Carnivory"
(coauthored with Natalia Vasey); Bruce Winterhalder, University of North Carolina, Chapel Hill, "Intragroup Resource
Transfers: Comparative Evidence, Models, and Implications for Human Evolution."

[Reference]
References Cited

[Reference]
ISAAC, G. L. 1978. The food-sharing behavior of proto-human hominids. Scientific American 238:go-io8.
WASHBURN, S. L., AND C. LANCASTER. 1968. "The evolution of hunting," in Man the hunter. Edited by R@ B. Lee and
1. DeVore, pp. 293-303. Chicago: Aldine.

[Author note]
CRAIG B. STANFORD AND HENRY T. BUNN Department of Anthropology, University of Southern California, Los
Angeles, Calif goo8q, U.S.A. (stanford@almaak.usc.edu)/Department of Anthropology, University of Wisconsin, Madison,
Wis. 537o6-1395, U.S.A. (htbunn@facstaff. wisc.edu).
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