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MEAT-EATING &
HUMAN EVOLUTION
THE HUMAN EVOLUTION SERIES

Editors
Russell Ciochon, University of Iowa
Bernard Wood, George Washington University

Editorial Advisory Board

Leslie Aiello, University College, London
Alison Brooks, George Washington University
Fred Grine, State University of New York, Stony Brook
Andrew Hill, Yale University
David Pilbeam, Harvard University
Yoel Rak, Tel-Aviv University
Mary Ellen Ruvolo, Harvard University
Henry Schwarcz, McMaster University

African Biogeography, Climate Change, and Human Evolution

edited by Timothy G. Bromage and Friedemann Schrenk

Meat-Eating and Human Evolution

edited by Craig B. Stanford and Henry T. Bunn
MEAT-EATING &
HUMAN EVOLUTION

EDITED BY
Craig B. Stanford & Henry T. Bunn

OXPORD
UNIVERSITY PRESS

2001
OXFORD
UNIVERSITY PRESS
Oxford New York
Athens Auckland Bangkok Bogota Bombay Buenos Aires
Calcutta Cape Town Dar es Salaam Delhi Florence Hong Kong
Istanbul Karachi Kuala Lumpur Madras Madrid Melbourne
Mexico City Nairobi Paris Shanghai Singapore Taipei Tokyo Toronto Warsaw
and associated companies in
Berlin Ibadan

Copyright © 2001 by Oxford University Press, Inc.

Published by Oxford University Press, Inc.
198 Madison Avenue, New York, New York 10016
Oxford is a registered trademark of Oxford University Press
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording, or otherwise,
without the prior permission of Oxford University Press.

Library of Congress Cataloging-in-Publication Data

Meat-eating and human evolution / edited by Craig B. Stanford and Henry T. Bunn.
p. cm.
Includes bibliographical references and index.
ISBN 0-19-513139-8
1. Prehistoric peoples—Food. 2. Fossil hominids. 3. Meat—History.
4. Human evolution. I. Stanford, Craig B. (Craig Britton), 1956–
II. Bunn, Henry T.
GN799.F6 b M43 2001
599.93'8—dc21 00-036745

9 8 7 6 5 4 3 2 1
Printed in the United States of America
on acid free paper
 C. S. dedicates the volume to his parents,
Jacqueline and Leland Stanford, Jr.

H. B. dedicates the volume to his family

and we both dedicate the book to the memory of Glynn Isaac

This page intentionally left blank
 Contents

Contributors xi

Introduction 3
Craig B. Stanford
Henry T. Bunn

I. Meat-Eating and the Fossil Record

1 Deconstructing the Serengeti 13

Martha Tappen

2 Taphonomy of the Swartkrans Hominid Postcrania and Its Bearing

on Issues of Meat-Eating and Fire Management 33
Travis R. Pickering

3 Neandertal Hunting and Meat-Processing in the Near East:

Evidence from Kebara Cave (Israel) 52
John D. Speth
Eitan Tchernov

4 Modeling the Edible Landscape 73

Jeanne Sept
viii Contents

II. Living Nonhuman Analogs for Meat-Eating

5 The Dog-Eat-Dog World of Carnivores: A Review of Past and

Present Carnivore Community Dynamics 101
Blaire Van Valkenburgh

6 A Comparison of Social Meat-Foraging by Chimpanzees and

Human Foragers 122
Craig B. Stanford

7 Meat and the Early Human Diet: Insights from Neotropical

Primate Studies 141
Lisa M. Rose

8 The Other Faunivory: Primate Insectivory and Early Human Diet 160
William C. McGrew

9 Meat-Eating by the Fourth African Ape 179

Margaret J. Schoeninger
Henry T. Bunn
Shawn Murray
Travis Pickering
Jim Moore

III. Modern Human Foragers

10 Hunting, Power Scavenging, and Butchering by Hadza Foragers and

by Plio-Pleistocene Homo 199
Henry T. Bunn

11 Is Meat the Hunter's Property?: Big Game, Ownership, and

Explanations of Hunting and Sharing 219
Kristen Hawkes

12 Specialized Meat-Eating in the Holocene: An Archaeological Case

from the Frigid Tropics of High-Altitude Peru 237
John W. Rick
Katherine M. Moore

13 Mutualistic Hunting 261

Michael S. Alvard

14 Intragroup Resource Transfers: Comparative Evidence, Models, and

Implications for Human Evolution 279
Bruce Winterhalder
 Contents ix

IV. Theoretical Considerations

15 The Evolutionary Consequences of Increased Carnivory

in Hominids 305
Robert A. Foley

16 Neonate Body Size and Hominid Carnivory 332

Natalia Vasey
Alan Walker

Conclusions: Research Trajectories on Hominid Meat-Eating 350

Henry T. Bunn
Craig B. Stanford

Index 361
This page intentionally left blank
 Contributors

Michael Alvard Jim Moore

Department of Anthropology Department of Anthropology
380 MFAC University of California, San Diego
State University of New York at Buffalo La Jolla, CA 92093
Buffalo, NY 14261
Katherine M. Moore
Henry Bunn American Section, Museum of Archaeology
Department of Anthropology and Anthropology
5240 Social Building University of Pennsylvania
Univ. of Wisconsin 33rd and Spruce Streets
Madison, WI 53706-1395 Philadelphia, PA 19104

Robert Foley Shawn Murray

Department of Biological Anthropology Department of Anthropology
University of Cambridge University of Wisconsin
Downing Street Madison, WI 53706-1395
Cambridge, CB2 3DZ
United Kingdom Travis Pickering
University of Witwatersrand Medical
Kristen Hawkes School
Department of Anthropology Department of Anatomical Sciences
University of Utah 7 York Road, Parktown 2193
Salt Lake City, UT 84112 Johannesburg, South Africa

William McGrew John Rick

Departments of Biology and Anthropology Department of Anthropology
Miami University Stanford University
Oxford, OH 45056 Stanford, CA 94305-2145

XI
xii Contributors

Lisa Rose Eitan Tchernov

Department of Anthropology Department of Evolution, Systematics,
Washington University and Ecology
Box 1114 Hebrew University-Jerusalem
St. Louis, MO 63130-4899 91904 Jerusalem
Israel
Margaret Schoeninger
Department of Anthropology Blaire van Valkenburgh
University of Wisconsin Department of Biology
Madison, Wl 53706-1395 U.C.L.A.
Los Angeles, CA 90095-1606
Jeanne Sept
Department of Anthropology Natalia Vasey
Indiana University Department of Anthropology
Bloomington, IN 47405 409 Carpenter Building
Penn State University
John Speth University Park, PA 16802
Museum of Anthropology
4009 Museums Building Alan Walker
University of Michigan Department of Anthropology
Ann Arbor, MI 48109 409 Carpenter Building
Penn State University
Craig Stanford University Park, PA 16802
Jane Goodall Research Center
Department of Anthropology Bruce Winterhalder
University of Southern California Department of Anthropology
Los Angeles, CA 90089-0032 CB #3115 Alumni Biding.
University of North Carolina
Martha Tappen Chapel Hill, NC 27599
Department of Anthropology
University of Minnesota
395 HHH Center
301 19th Ave. South
Minneapolis, MN 55455
MEAT-EATING &
HUMAN EVOLUTION
This page intentionally left blank
 Introduction

Craig B. Stanford
Henry T. Bunn

M;I ore than 30 years after the publication of Man the Hunter, the
role of meat in the early human diet remains a central topic of human
evolutionary research. There is little doubt that meat-eating became increasingly
important in human ancestry, despite the lack of direct evidence in the fossil record
of how meat was obtained, or how much was eaten, or how often, or how exactly
increasing importance of meat-eating may have contributed to the rise of the genus
Homo. Although the fossil evidence is becoming clearer on these issues, we still
lack key evidence about early hominid behavioral ecology. Information about meat-
eating patterns from modern nonhuman primates, from modern foraging people,
and from the fossil record could all contribute to a clearer picture of early human-
ity than we have at present.
With this goal in mind, a workshop was held October 2-5, 1998, on the campus of
the University of Wisconsin, Madison. "The Early Human Diet: The Role of Meat,"
sponsored by the Wenner-Gren Foundation for Anthropological Research, brought
together 18 participants representing several subfields of human origins research.
Papers were presented at the workshop by Michael Alvard, Henry Bunn, Robert Foley,
Kristen Hawkes, William McGrew, Katharine Milton, Travis Pickering, John Rick,
Lisa Rose, Margaret Schoeninger, Jeanne Sept, John Speth, Craig Stanford, Mary Stiner,
Martha Tappen, Blaire Van Valkenburgh, Alan Walker, and Bruce Winterhalder.
Why publish a volume on meat-eating at this time? Despite its importance in the
evolutionary ecology of the Hominidae, scholars from different disciplines have
only rarely gathered to discuss the topic. Few of the contributors to this volume
had sat in the same group to discuss the crosscutting aspects of their work before
the Madison workshop. Most of the participants work in the field of biological
anthropology or archaeology; lack of intellectual crossfertilization may simply re-
flect increasing specialization within the discipline.

3
4 Introduction

Each era in the study of human behavioral origins has treated meat-eating in its
own way, based on the most reasonable interpretations of the available data. Since
Raymond Dart (1953), reconstructions of early hominid behavior have revolved
around dietary issues, due to the recognition that among many social animals in-
cluding nonhuman primates, social behavior and grouping patterns are profoundly
influenced by the need to balance energy output with nutrient energy intake. The
diet of most higher primates consists largely of leaves and fruit, and foraging for
these consumes most of each day. Including a highly concentrated packet of nutri-
ents and calories, such as meat represents, may have provided emerging humans
with a key nutritional supplement that favored the evolution of other key traits, such
as cognition.
From the 1960s until the early 1980s, consideration of meat-eating generally
focused on the importance of hunting to early human social patterns (Washburn
and Lancaster 1968; Tiger and Fox 1971; Suzuki 1975; Lovejoy 1981; Hill 1982;
Tooby and DeVore 1987). In this earlier era, the most influential and ultimately
infamous body of theory related to meat-eating was Man the Hunter. The idea that
hunting was the seminal behavior accounting for the expansion of the human brain
neocortex and higher intelligence emerged from a conference of the same name
held in April, 1966, in Chicago. About 75 scholars gathered to discuss the behavior
and status of foraging people ("hunter-gatherers") in the world at that time. The
volume that followed, edited by Richard Lee and Irven DeVore, included a chapter
by Sherwood Washburn and Chet Lancaster entitled "The Evolution of Hunting,"
in which Washburn and Lancaster hypothesized that hunting was among the most
fundamental of human behavioral adaptations. They proposed that the importance
of communicating and coordinating big game hunting placed a premium on intel-
ligence and the expansion of the brain's neocortex. Because hunting is primarily a
male activity in modern, and presumably ancient, human societies, this would have
accounted for the large size of the human brain in males. By ignoring the role of
females in the evolution of human brain size, Washburn and Lancaster unleashed a
firestorm of criticism. Some anthropologists (e.g., Tanner and Zihlmann 1976) took
issue with the assumption that meat composed a substantial or important portion of
the early human diet. Others argued that a predatory view of human cognitive ori-
gins was rooted in male-biased science. Ironically, the consensus of opinion at the
Man the Hunter conference was that meat is of relatively little nutritional impor-
tance in the diets of the same modern tropical foragers.
The legacy of Man the Hunter was long-lasting in academic discussions of meat-
eating and human origins. The debate may have even accounted in part for the rise
of feminist theory in anthropology in the 1970s (Stanford 1999). The backlash
against Man the Hunter led many anthropologists to reject hunting as an important
subsistence mode among early hominids and led others to reject meat-eating as an
important part of the early human diet.
Beginning in the 1980s, the hunting paradigm fell victim to reinterpretations of
archaeological sites, which suggested that the cooperative, predatory tendencies of
early humans had been misinterpreted. Data from Plio-Pleistocene sites were in-
creasingly interpreted as evidence of meat procurement by scavenging rather than
by hunting (Binford 1981; Isaac and Crader 1981; Shipman 1986; Blumenschine
 Introduction 5

1987; Potts 1988). Bunn (1982) and Bunn and Kroll (1986) advocated both scav-
enging of large ungulate carcasses and hunting of smaller prey based on their analysis
of Plio-Pleistocene material from Olduvai Gorge. Some early advocates of the
importance of scavenging (Howell 1968; Schaller and Lowther 1969) had pro-
posed this behavior as an adjunct to hunting, but for the more recent work scav-
enging was often proposed as the primary or even sole means of carcass acquisi-
tion. Shipman and Potts (1981) and Shipman (1986) showed that some Pliocene
bone assemblages had unambiguous evidence of hominid cutmarks made on top
of carnivore toothmarks, supporting a scavenging foraging mode for early genus
Homo. Binford (1981) took the most extreme view, rejecting the possibility that
any taxon of hominid prior to earliest Homo sapiens would have had the cognitive
capacity for cooperative hunting or food-sharing. Isaac's seminal (1978) work on
food acquisition and food-sharing among early hominids was part of a movement
to consider the Pliocene past by use of analogy with the better-understood present.
During the workshop we returned time and again to Isaac's ideas and agreed that
his food-sharing model, put aside during the rush to "dehumanize" early hominids
during the 1980s, accords as well with field data today as it did then.
In the 1990s, a more balanced view of hunting and scavenging has prevailed,
which this volume attempts to represent. The current perspective has been based
on research in the three areas covered by this volume: meat-eating by nonhuman
primates and their analogs, meat-eating by modern foragers, and evidence of meat-
eating in the fossil record. There is a growing consensus among researchers study-
ing the fossil record that earlier dichotomies between hunting and scavenging were
simplistic and ultimately false. This perspective was evident at the Wenner-Gren
workshop, in which the long-standing debate over the occurrence and importance
of hunting and scavenging by early hominids was rarely at issue. There is a recog-
nition today that this dichotomy has eroded with the collection of data from a vari-
ety of research sites. Thus, Blumenschine's (1986) argument for an exclusive
scavenging niche based on a reconstruction of the ecology of Pliocene Serengeti
was extremely valuable, but no longer accords well with data from reconstructions
of hominid behavioral ecology from other habitats (e.g., Tappen, this volume). There
are no obligate scavengers among living mammals; carnivores from lions to hy-
enas typically acquire meat by either hunting or scavenging as the opportunity arises.
Bunn and Ezzo (1993) and Bunn (this volume) argue for a mode of hominid sub-
sistence based on the opportunistic hunting and pirating ("power scavenging") of
large mammalian carcasses in a manner that resembles what many carnivores do
today. This does not mean that passive scavenging might not have been important
in some periods and among some taxa in hominid evolution; only that the strict
hunting versus scavenging debate of the 1980s seems to have given way to a more
realistically complex view of Pliocene hominid behavioral ecology.

Approaches to the Study of Meat-Eating

Some definitions are in order before we proceed further with a discussion of meat-
eating. First, by meat-eating we refer to the consumption of vertebrate fauna (but
6 Introduction

see McGrew, this volume, for invertebrate faunivory), including muscle, viscera,
the skeleton, and associated body tissues. "Meat" is thus more properly referred to
as "carcass biomass," but for purposes of this volume it is understood that meat-
eating encompasses all body tissues. The nutrient and caloric values of mamma-
lian carcasses have been studied by a range of scholars in fields ranging from bio-
chemistry and nutritional sciences to archaeology, and for an equal variety of
reasons. This volume contains a number of chapters that discuss the nutrient and
caloric properties of meat but none that examines in detail the biochemical basis
for meat as a valuable nutrient source (that is, the amino acids, fats, etc., contained
in a carcass). This is perhaps a necessary failing in that all the chapters herein ac-
cept the (admittedly incomplete) received wisdom about why carnivores and om-
nivores live on diets that are partially or wholly the meat of other animals.
We include both scavenged carcasses and hunted live prey when discussing meat-
eating as a dietary/behavioral adaptation. Considering these as separate foraging
modes makes sense, even though there are no living mammals that do one without
at least sometimes also doing the other.
The lines of evidence that were presented at the conference encompassed the
three fields below, with many chapters crosscutting two or more of these. In addi-
tion, two theoretical issues directly related to meat-eating in human evolution were
included that did not fall neatly into any of the three areas below.

Meat-Eating by Nonhuman Analogs

Recent field data on hunting behavior by wild chimpanzees, building on Teleki (1973)
and Goodall's (1986) work, have shown that chimpanzees consume more meat, at
least at some study sites, than previously thought (Stanford 1996, 1998). At some
sites, chimpanzees hunt cooperatively (Boesch and Boesch 1989, Boesch 1994). The
level of predatory cooperation seen among wild chimpanzees refutes Binford's
argument that early hominids would not have been cognitively able to engage in
cooperative hunting and food-sharing. In addition, the growing realization that chim-
panzee populations display cultural diversity paralleling that of the most technologi-
cally simple human societies (McGrew 1992) provides much insight into the likely
cultural aspects of early human technologies and other behaviors.
Using nonhuman primates to interpret the meat-eating behavior of our ances-
tors is, however, fraught with problems. Chimpanzees and modern humans share
an ancestor that lived some six million years ago; we cannot assume that modern
chimpanzees are very similar to the ancestral chimpanzee any more than we would
think that modern people are very much like the ancestral hominid. The chimpan-
zee dietary adaptation reflects life in a wide variety of habitat types, some of which
may never have been inhabited by Pliocene hominids. Moreover, among the four
great apes, only the chimpanzee is an avid hunter and consumer of meat. Using the
chimpanzee as a presumptive model of meat-eating patterns in an early australo-
pithecine necessarily ignores other living exemplars, like the bonobo, that offer a
contrasting view.
Nevertheless, chimpanzees are valuable referential models of early human be-
havioral ecology because they enable us to go beyond the one-dimensional portrait
 Introduction 7

that the fossil record provides of extinct taxa. Because it is in most cases extremely
fragmentary, the fossil record can deceive us into accepting a single, well-documented
site as representative of a species' biology. Chimpanzee behavioral diversity across
wide geographic areas, due to both ecological influences and local cultural tra-
ditions, offers an important lesson for students of early hominid behavioral ecol-
ogy. Chimpanzee behaviors from tool use to hunting techniques to grooming styles
vary from population to population. Likewise, we should expect that a species of
Australopithecus or early Homo may have been an avid scavenger of large carcasses
at one site and an avid hunter but not a scavenger at a contemporaneous site 100 km
away. Chapters in this volume by `McGrew, Rose, and Schoeninger et al. present
ideas and data related to the consumption of meat by nonhuman primates.
We need not limit ourselves to primates when attempting to reconstruct the be-
havior and ecology of the earliest hominids. Van Valkenburgh reconstructs Pliocene
African ecosystems in which early Homo would have been one component and,
using data on diet and body weight, argues that feeding competition from other,
larger, meat-eating species would have been major factors in the behavioral ecol-
ogy of these taxa.

Meat-Eating by Modem Foraging People

Field studies of modern foraging societies have done much to show how and why
they obtain meat. Meat may compose only a small part of the diet, but the compo-
sition of the overall diet and its seasonal variation provide opportunities for hypoth-
esis testing. Likewise, the ways in which prey are caught, or carcasses scavenged,
and then butchered and distributed to group members are still poorly understood
for many foraging societies. Studies of tropical and subtropical foragers, especially
of the behavioral ecology of the Hadza in East Africa (Hawkes et al. 1991), the
IKung in southwestern Africa (Lee 1979), the Efe in eastern Congo (Bailey and
Peacock 1988), and the neotropical Ache of Paraguay (Hill and Hawkes 1983;
Kaplan and Hill 1985), have tested hypotheses about the pattern and purpose of
meat-eating. These studies have shown that elements of the behavioral ecology of
modern people, such as nutrient/caloric costs and benefits of foraging for plant versus
animal foods (Hawkes 1993) and the pattern and sequence of carcass transport and
consumption (O'Connell and Hawkes 1988), provide appropriate and valuable
comparisons with analogous behaviors among other living primates having similar
energetic exigencies.
There is a long-standing debate about the utility of modern foragers in studying
human evolution; some scholars assert that studying modern people with an eye
toward the past is inherently useless and possibly even racist. Humans living with
relatively simple technologies, who forage for a living from their forest or grass-
land environment, make decisions every day about which foods to forage for and
which to pass by, or about which parts of an animal carcass they will relish and
which they will discard. These decisions, no matter how culturally influenced, are
tied to the nutritional health and reproductive lives of the men and women in the
group. As such they can be examined, and questions can be asked about the deci-
sions themselves. There is no doubt that even among the most remote foraging
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