You are on page 1of 17


Johns Hopkins University

Scavenging or Hunting in Early Hominids:

Theoretical Framework and Tests
Evidence from Bed I, Olduvai, supports the hypothesis that scavenging, not hunting, was the
major meat-procurement strategy of hominids between 2 and 1.7 million years ago. Data used to
evaluate the hunting and scavenging hypotheses are derivedjom studring cut marks on Bed I
bovids, comparing adaptations necessaryfor scavenging with those of ear& hominids, and a paleoecological reconstruction of Bed I carcass biomass, carnivore guild, and hominidforaging area.

to that of
modern hunter-gatherers. Whether carrying (Hewes 1961, 1964) tool making (Darwin 1871; Washburn 1960), food sharing (Isaac 1978, 1983), or seed eating (Jolly 1970)
is seen as the crucial adaptation in hominid evolution, hunting and meat eating are often
given a major place in early hominid life (Ardrey 1961, 1976; Bunn 1982, 1983b; Hill
1982; Isaac and Crader 1981; Tiger and Fox 1971; Washburn and Lancaster 1968; Washburn 1978). Although scavenging has been suggested, often as a behavioral transition
between foraging for plant foods and hunting, it has only recently begun to be evaluated
critically as a distinct and important adaptation (Binford 1981, 1984;Dunbar 1983; Isaac
and Crader 1981; Isaac 1983; Potts 1982, 1984; Szalay 1975).
Previously, I presented preliminary evidence that hunting accompanied by systematic
butchery and food sharing, as is typical of modern hunter-gatherers, is not supported by
evidence from Bed I, Olduvai Gorge, Tanzania (Shipman 1984,1981; Potts and Shipman
1981; Shipman 1983). Bed I sites are appropriate for testing hypotheses about early hominid food-procurement strategies because of their antiquity (2-1.7 million years, Hay
1976), their numerous, well-preserved faunal remains associated with artifacts, and the
well-documented geology, archeology, and taxonomy.
In this paper, I articulate and test hunting and scavenging hypotheses, using scanning
electron microscope studies of cut marks and carnivore tooth marks on Bed I fossils and
ecological, physiological, and behavioral studies of modern hominids and scavengers. Because it is unclear which hominid species preserved in Bed I manufactured the Oldowan
tool industry (Leakey 1971), I use the term Oldowan to refer to whichever hominid(s)
were responsible for these activities.

The Hypotheses: Hunting versus Scavenging

The hunting hypothesis postulates that a major factor in Oldowan life was the development of hunting techniques and tools that fostered the development of central places
or base camps; these served as temporary dwelling places and rendezvous at which animal foods were shared with or supplied to other members of the social group. Variants
of this hypothesis are expressed in many works that emphasize the role of hunting and its
correlated behaviors in human evolution, although scavenging is often mentioned briefly.
As a heuristic device designed to clarify hypothesis evaluation, the hunting hypothesis
is Assistant Prgessor,Department of Cell Biology and Anatomy, School of Mcdicinc,Johns Hopkins University,
Baltimore, M D 21205.




[88, 1986

articulated here excludes any significant contribution to the diet by scavenging. Hunting is defined as the intentional killing of animals larger than 5 kg by hominids using
tools and weapons. The contribution of meat to the total diet obtained by hunting is unquantified but significant (Isaac 1983:13), the rest of the diet being provided by gathered plant foods.
Behaviors closely associated with hunting, such as butchery and systematic disarticulation to permit food transport and sharing, are more readily identified in the fossil
record than hunting per se. I explicitly assume here that Oldowan hunters engaged in
these carcass-processing behaviors. It is theoretically possible that Oldowans hunted
without engaging in any of these ancillary behaviors, in which case hunting would be
nearly invisible archeologically or paleontologically. Propositions for testing the hunting
hypothesis were generated from the extensive ethnographic literature about modern
hunter-gatherers (e.g., Bicchieri 1972; Binford 1981; Coon 1971; Gifford 1977; Gould
1968, 1980; Lee and DeVore 1968, and references therein; Marks 1976; Marshall 1965;
Service 1979; Turnbull 1965a, 196513; Winterhalder and Smith 1981, and references
therein; Yellen 1977):
(1) If prey animals were disarticulated, then verified cut marks will occur near joints
in frequencies (about 90%) comparable to those observed on more recent, butchered
(i.e., disarticulated) prey remains.
(2) If both skin and meat were removed from bones, then skinning marks will be substantially more frequent than meat-removal marks (75% vs. 25% respectively), because
the skin is separated from distal limb elements overlaid by little flesh. Carnivores interested primarily in obtaining meat produce a complementary pattern ofdamage, with over
75% of tooth marks occurring on meat-bearing bones (personal observation).
(3) If Oldowans were primarily hunters, then usually their cut marks will be overlaid
by carnivore tooth marks, when sets of overlapping marks of different origin are found.
A quantitative prediction about the frequency of overlapping cut marks and tooth marks
cannot be made because there are no comparable data for assemblages known to have
been hunted by stone-tool-wielding hominids and then scavenged.
The scavenging hypothesis proposes that the Oldowans were poor hunters, infrequently capable of killing and defending their own prey. Instead, Oldowans relied mostly
on scavenging to obtain meat, skin, or other substances from carcasses. Scavenging supplemented plant food foraging and did not provide the major portion of dietary intake,
since such a situation is unknown among living mammals (Houston 1979). The contribution of scavenging to the diet is set at 33%, a figure chosen to indicate that scavenging
is as significant a food-procurement strategy to Oldowans as it is to the most successful
mammalian scavenger today, the spotted hyena. Scavenging refers both to obtaining
meat or other substances from carcasses killed by other species and to carrion eating, or
consuming partial or whole animals dead of nonpredatory causes.
Ethnographic data are lacking on predominantly scavenging peoples, so predictions
for testing the scavenging hypothesis were generated by assuming, first, that the carcassprocessing patterns typical of hunter-gatherers would be absent, and second, that scavenging, nonprimate species serve as appropriate analogues to a limited extent:
(1) If scavenging Oldowans access to carcasses was brief, then the clustering of cut
marks near joints, indicative of systematic disarticulation, will be absent; the location of
cut marks will closely resemble that of carnivore tooth marks.
(2) Since scavengers access to the choicest body parts are denied by the primary predators, Oldowans processing activities will occur more frequently (>25%) on non-meatbearing areas and less frequently (< 75%) on meat-bearing areas than those of carnivores.
(3) If hominids frequently scavenged from carnivore kills, then their cut marks will
commonly overlie tooth marks when sets of overlapping marks are found.
(4) If scavenging was an important Oldowan behavior, then the physical or behavioral




adaptations that distinguish modern scavenging species from predominantly hunting

ones will be present among the Oldowans.
(5) If Oldowans practiced a scavenging and foraging lifestyle, then reconstruction of
the Bed I, Olduvai, ecosystem will reveal sufficient herbivore biomass and predatory carnivores to provide carcasses for a scavenging hominid.
Binford (1984), Potts ( 1982), and Vrba (1980) suggest that scavenged assemblages
show characteristic patterns of skeletal and age class representation. These criteria are
not used because some studies (Bunn 1982; Shipman, Davis and Bosler, unpublished
data) of scavenged assemblages do not show the suggested skeletal and age class representation patterns. In addition, Bed I assemblages show signs of a complex taphonomic
history involving multiple agents (Potts and Shipman 1981) over at least several years
(Potts 1982, 1984), which may have confused or obscured patterning created by any one

Materials and Methods

A comprehensive survey of damage on major limb bones of bovids ( N > 2,500 specimens) from Bed I was undertaken. Bovids were selected as the most numerous and most
probable prey species for any predator. Limb bones were utilized because they are abundant, well preserved, frequently identifiable to taxon, and often cut-marked at recent
butchery sites (Bunn 1983a; Crader 1983; Guilday et al. 1962). All available bovid limb
bones were inspected by eye and Iight microscope. All marks suspected ofbeing cut marks
were replicated, as were a random sample of suspected toothmarks. All fossils identified
as cut-marked by previous researchers were also replicated as were those bearing overlapping sets of marks, regardless of element or taxon. One to three areas on each of a total
of 203 fossils were replicated and inspected using standard procedures (Rose 1983; Shipman and Rose 1983a, 1983b). SEM inspection verified the identity of cut marks on 76
bones and carnivore tooth marks on 70 bones; many anatomical, preparators, and rodent
gnawing marks were also identified.
Criticism (Bunn 1982, 1983b) that replication and SEM-inspection are unnecessary
spurred a test of the accuracy of more gross means of mark identification. I reevaluated
230 marks identified by eye or light microscopy by Bunn, Potts, or Shipman as cut marks,
without knowledge of the original identifications, using the SEM. Only 55%-60% of
these showed the microscopic criteria diagnostic of cut marks. There was no significant
variability in different observers success rates. It is highly unlikely that the SEM identifications are incorrect, since every known cut mark in a large control sample ( N > 1,000)
inspected by SEM shows these microscopic features. The only taphonomic circumstance
that produces marks microscopically identical to cut marks-trampling of bones in caves
on top of sharp-edged rockfalls (Oliver 1984)-did not occur at Olduvai. Tooth marks,
vascular grooves, and preparators marks were most commonly misidentified as cutmarks; rodent gnawing and heavy carnivore chewing are often readily identified at a gross
It is suggested that SEM use is not generally warranted if (1) the assemblage was created or modified by Homo sapiens, a species of known habits and (2) the observed patterning of presumed cut marks conforms to ethnographic reports for that region. In one such
case (Villa and Shipman, unpublished data) there was over 90% concordance in identifications made by eye/light microscopy and SEM. However, more extensive SEM use
is warranted in any study that relies on the sequence in which sets of overlapping marks
were made.
Statistical evaluation of the Olduvai data require a control assemblage of bones hunted
and butchered by hominids. Quantitative data for large, ethnographically documented
assemblages with appropriate species are lacking; data from a Neolithic carcass-processing site (Prolonged Drift, Gifford et al. 1981) were used instead. Gross mark identifications by Gifford et al. (1981) were accepted, since criteria suggested above were met;



[88, 1986

further, the assemblage lacks evidence of carnivore activity. Using data from Prolonged
Drift, rather than a modern site, offers advantages. Prolonged Drift preserves stone tool
cut marks rather than metal tool marks, which are more abundant (personal observation). Alsa, Gifford et al. (1981) provide detailed quantitative data on 296 cut marks on
bovids similar to those at Olduvai, out of an identifiable subset (N = 3,700 bones) of the
total assemblage (N > 20,000); these data are sufficient to establish clear patterns of carcass utilization. Finally, the interpretation of Prolonged Drift has met with little or no
Verified cut marks and tooth marks were classified as occurring in near joint or midshaft locations; when data were insufficient to determine which of these categories was
appropriate, marks were assumed to occur in a near joint location (thus favoring the
hunting hypothesis). Verified marks were classified as being located on meat-bearing
bones (the proximal elements of each limb) or non-meat-bearing bones (the elements
distal to the femur or humerus). Forty-two sets of overlapping marks were located and
replicated, Only 13 sets included both cut marks and carnivore tooth marks and showed
clear indications of the sequence in which the marks were made (Shipman and Rose
Literature on modern African ecosystems was used to identify adaptations and ecological prerequisites to successful scavenging, as distinct from hunting. Primary data on African animals are derived in general from Schaller (1968, 1972), Kruuk (1972, 1976),
Bertram (1973, 1975), Estes and Goddard (1967), Frame and Frame (1976), Rudnai
(1973), and Van Lawick-Goodall and Van Lawick (1970). Useful reviews and analyses
are found in Bertram (1979), Curio (1976), Eaton (1979), Ewer (1973), Houston (1979),
Nowak and Paradiso (1983), and C. Pennycuick (1971, 1979). For ease of discussion below, references will be cited only for specific data on attributes of carnivore and avian
In addition, data on Olduvai ecology and fauna, on various metabolic relationships,
and on Oldowan population densities were used in reconstructions. All estimates and
calculations were based as firmly as possible on documented information and established
energetic and ecological principles, but the results may not be accurate much past the
order of magnitude. All estimates were made conservatively, to test the scavenging hypothesis stringently. In addition, it was established a priori that the prediction of feasibility would be upheld only if available carcass biomass generously exceeded consumption.

Results and Discussion: Cut Mark Studies

The first prediction of each hypothesis depends upon the distribution of cut marks on
the Olduvai fossils in terms of near joint and midshaft locations. Chi-square tests reveal
that the Olduvai cut marks are distributed significantly differently from the Prolonged
Drift cut marks (Table 1) and do not cluster near joints as predicted by the hunting hypothesis. In contrast, the Olduvai cut mark distribution is indistinguishable from that of
the 70 carnivore tooth marks, as predicted by the scavenging hypothesis. These results
are not significantly altered by the exclusion ofcut marks ofambiguous location (assumed
to be near joint) from the sample.
The second predictions of the two hypotheses were both fulfilled (p < 0.05 in each case:
Table 2) but are not mutually exclusive. The results could be interpreted equally validly
to mean either that hominids utilized carcasses opportunistically (scavenged) or that they
utilized carcasses fully and systematically (hunted).
Of the 13 overlapping sets of marks, 8 showed that the carnivore tooth marks were
made first and the cut marks were made second; in the remaining 5 cases, the situation
was reversed. These data are direct, if limited, evidence that Oldowans scavenged from
carnivore kills on occasion. These results do not differ statistically from a hypothetical
situation in which 6.5 cut marks overlie tooth marks and 6.5 tooth marks overlie cut




Table 1
Distribution of cut marks and tooth marks on Olduvai bovids relative to joints compared
with that of cut marks from Prolonged Drift.

(a) Olduvai cut marks

Near joint






(b) Prolonged Drift

(c) Olduvai tooth marks

avs. b: chiz = 86.3
1 degree of freedom
p c 0.001*
b vs. c: chiz = 125.0
1 degree of freedom
p c 0.001*
a vs. c: chiz = 0.143
1 degree of freedom
p > 0.05

*Indicates level of probability accepted as significant.

Table 2
Distribution of cut marks and tooth marks on Olduvai bovids relative to major muscle
masses compared with that at Prolonged Drift.
Meat-bearing Non-meatbones
bearing bones
avs. b: chiz = 2.5
(a) Olduvai cut marks
1 degree of freedom
p > 0.05
(b) Prolonged Drift
b vs. c: chi2 = 50.9
1 degree of freedom
p C 0.005*
a vs. c: chiz = 17.9
(c) Olduvai tooth marks
1 degree of freedom
p C 0.005*


*Indicates level of probability accepted as significant.

marks. These results do differ significantly (chi-square = 4.4, 1 d.f., p C 0.05) from a
hypothetical situation in which all 13 tooth marks overlie cut marks, the extreme case
expected if Oldowans only hunted.
In general, the cut mark data support the scavenging hypothesis and do not support
the hunting hypothesis as stated here. A possibility that cannot be eliminated is that
hunting was carried out by Oldowans but that neither disarticulation nor transport or
sharing of food occurred. I can see no means to transform this possibility into a testable
hypothesis at present.

Results and Discussion: Adaptations to Scavenging

The Serengeti Plains ecosystem, adjoining Olduvai Gorge, is inhabited by nine species
of medium- and large-sized carnivores: the lion, Panthera Leo; the spotted hyena, Crocuta
crocuta; the leopard, Panthera pardus; the cheetah, Acinonyx jubatur; the striped hyena,
Hyaena hyuena; the hunting dog, Lycaonpictus; and three jackals, Canis mesomelas, C.adustus,
and C. aureus. Differences among the three jackals are poorly documented, so they are
treated as a single species (Canis sp.) here. Only the cheetah and the hunting dog are
almost never observed to scavenge. Each of the others scavenges to a greater or lesser
extent (up to about 33%; Bertram 1979:222-223). In general, when the migratory herbivores are absent, these part-time scavenging species turn to alternative food sources
(fruit, hunted prey); when the migratory herds are present, carcasses are abundant and




these species scavenge more frequently. In short, they scavenge when they can and hunt
or forage when they must. This flexibility suggests that hunting and scavenging lie along
a behavioral continuum; nonetheless, there are powerful adaptive differences between
those that do and do not scavenge. Since no living mammal scavenges for all of its food,
I discuss the only purely scavenging vertebrates in Africa today: species of raptors, referred to here collectively as vultures, that are obligate or nearly exclusive scavengers.
For analysis, species were divided into three categories that represent a gradient from
hunting to scavenging: (1) predatory carnivores (cheetah, hunting dog) that rarely scavenge; (2) scavenging carnivores (jackal, striped and spotted hyenas, leopard, lion) that
scavenge part-time; (3) obligate or exclusive scavengers (the vultures). Features typical
of each group were sought from the literature, Houston (1979) being especially useful.
The major adaptation enabling the vultures to be exclusive scavengers is their energetically inexpensive mode of locomotion: flying. Since carcasses are always less abundant than potential prey, only a species that can cover large areas cheaply is assured of
finding enough c:arcasses. Although the cost of locomotion is important to scavengers, its
speed is not. Predatory carnivores are consistently faster, in some sense, than scavenging
ones. Vultures lessen the energetic costs oflocomotion, but also their speed, by gliding at
the prevailing wind speed. In short, speed is a trade-off for cost and endurance; the former
is more important to predators or hunters, the latter to scavengers.
Perhaps the second most important requisite of a successful scavenger is an adaptation
for locating carcasses. In vultures, it is a secondary benefit of their mode of locomotion,
which improves their vantage point, combined with keen eyesight. The most successful
mammalian scavenger, the spotted hyena, responds more rapidly to the sight of circling
vultures than do other carnivores (Kruuk 1972) and has a keen sense of smell (Ewer
The third adaptation is a means of dealing with interference competition over carcasses. Often, primary predators are forced either to defend a carcass or to relinquish it
to scavengers. Large body size or sociality are important determinants of success in interference competition (Eaton 1979); small-bodied scavengers usually specialize in
stealth and avoidance behaviors. The two strategies can be simply characterized as being
either a bully or a sneak. Predatory carnivores characteristics contrast sharply with
those of scavengers. Both cheetahs and hunting dogs are of intermediate size, and both
frequently lose interference encounters. Further, hunting dogs are so highly social that it
is hard to scavenge sufficient food for the group. Predatory carnivores maximize the meat
obtained per unit energy expended in a chase by adapting for speed in pursuit and in
carcass processing; scavenging carnivores maximize either the number of carcasses they
can appropriate (the bully strategy) or the amount of meat they can scavenge without
risking appropriation (the sneak strategy).
The fourth adaptation to scavenging is utilizing a reliable, alternative food source.
Larger scavenging carnivores hunt when scavenging fails; smaller ones rely on fruit and/
or insects for the bulk of their diet.
Finally, many living scavengers apparently possess physiological or behavioral adaptations for dealing with rotten food, but these would be undetectable in the fossil record.
The hominid fossil record from Bed I was examined to see if the four detectable adaptations were present. All of the Bed I hominids possess clear and unmistakeable adaptations for bipedalism (Johanson and White 1979;Johanson et al. 1982; Leakey and Hay
1979; Lovejoy et al. 1973; Lovejoy 1974; McHenry 1978, 1982; McHenry and Temerin
1979; Robinson 1972; Stern and Susman 1983; Susman and Stern 1982; Zihlman 1978;
Zihlman and Brunker 1979). It has often been overlooked that, at speeds of 1.1 to 1.7 m/
sec, bipedal walking is empirically at least as efficient as quadrupedal walking, if body
size, speed, and distance are held constant (Fedak and Seeherman 1979; Taylor et al.
1982; Taylor 1977 and personal communication). Bipedal walking is also more efficient
than quadrupedal walking as practiced by modern pongids (Rodman and McHenry
1980). Thus, bipedal walking fulfills the locomotor needs of a scavenger.




Evidence for efficient carcass location is also seen. Bipedalism inevitably raised the
horninids head and markedly improved its ability to spot items on the ground, such as
carcasses (e.g., Dart 1959; Howells 1959). Arguments that australopithecines and Homo
habilis are adapted to tree climbing and arboreality in addition to bipedalism (McHenry
1978; Prost 1980; M. D. Rose 1984; Senut 1981; Stern and Susman 1983; Susman and
Stern 1979, 1982; Vrba 1979) suggest an additional adaptation for improving vantage
To reconstruct strategies for dealing with interference competition, knowledge of body
size is needed. Estimates for Oldowans range from about 18 kg to 70 kg (Brain 1981;
Cronin et al. 1981; Holloway 1978; McHenry 1976; Steudel 1980). I take 35 kg to represent mean Oldowan body size here, since this figure occurs within the separate ranges
for each hominid species. All modern scavenging carnivores of comparable size or smaller
use a sneak strategy. Retaining arboreal adaptations enabled Oldowans to lessen competition further by retreating into the trees to consume scavenged bits (see Brain 1970,
1981). In addition, the use of stone tools can be viewed as a direct adaptation to speedy
removal of substances from carcasses. Thus, two adaptations suitable for a sneak scavenger were present. Scavenging in social groups is also possible, but the cut-mark data
indicate this occurred rarely, briefly, or not at all. Binford (1984) suggests temporal strategies might lessen competition with largely crepuscular or nocturnal carnivores; this
plausible strategy is, unfortunately, difficult to test with evidence from the fossil record.
The most probable alternative food source for Oldowans was fruit, judging from dental
microwear data (Walker personal communication; Walker 1980, 1981). This pattern of
scavenging and frugivory is documented for striped hyenas, which are of comparable
body size to Oldowans (Kruuk 1976). Since the locomotor needs of scavenging are the
same as those of foraging for unpredictably distributed, stationary resources, such as
fruit, an individual can easily forage and scavenge simultaneously.
In summary, all adaptations ofscavengers likely to be observed in the fossil record were
present in Oldowans.

Results and Discussion: Ecological Feasibility

Was the biomass in ancient Olduvai suficient to make scavenging by Oldowans a feasible strategy?
The first consideration is that the Bed I carnivore guild was larger than todays. In
addition to rough equivalents of the modern Serengeti carnivores, Bed I had two extra
species of large, sabre-toothed cats, notable for their highly specialized, slicing teeth
(Walker 1984). These were highly predatory carcass providers adapted for meat eating,
not scavenging (Ewer 1973; Savage 1978); they were formidable opponents in interference competition (Brain 1981; Van Valkenburgh personal communication).
However, herbivore biomass was also larger. Faunal, geologic, isotopic and palynologic studies (Gentry and Gentry 1978a, 1978b; Hay 1976; Ceding et al. 1977; Bonnefille
1977 [cited in Potts 19821 ) all suggest Bed I rainfall exceeded current levels (800 mm/
yr; Norton-Griffiths et al. 1975) by 100-200 mm/yr. Since herbivore biomass in Africa is
a function of annual rainfall (Coe et al. 1976), it can be estimated that rainfall of 9001,000 mm/yr in the Serengeti would produce a 100%-400/~increase in prey biomass
over current levels if the habitat remained relatively open. Open habitat was likely, since
the Serengeti soils inhibit tree growth both now and in Bed I times (Sinclair 1979;sHay
1976). A realistic estimate of Bed I herbivore biomass is more than that of the Serengeti
today, or about 12,300 kg/km2 (East 1984). This reconstructed density is surpassed in
the modern Serengeti during the rainy season (L. Pennycuick 1975) and is well within
the realm of possibility.
The annual Bed I carcass biomass, expressed here as kilograms of dead herbivores
(kilocarcasses or kg-c) per square kilometer, is estimated using the average annual mortality rate of 16% across all ungulate species (Houston 1979) for the Serengeti today. A



[88, 1986

similar mortality rate in Bed I times would yield 1,968 kg-c/km2per year. How many kgc/km2 is an Oldowan likely to discover?
Foraging radius (r) is defined as the maximum distance that can be traveled by an
animal from a food source to another point (base camp, den, sleeping tree, waterhole,
tool cache, etc.) without undergoing net energy loss on the trip back or onward to a food
source. C. Pennycuicks basic equation (1979:167) is modified to allow 12 hours daily for
foraging, since Oldowans were almost certainly diurnal.

4 Em 4kV
the energy extracted from a full gut load
V = velocity of travel in meters per second
Em = basal metabolic rate (a function of m, or body size)
k = constant representing the energy required to propel an animal a unit distance, based on the metabolic cost of locomotion (Elz) in addition to Em.
Oldowan body weight is set at 35 kg. e varies between about 82,000 joules/kg of consumers body weight for grass eaters to at least 328,000 joules/kg for meat eaters; fruit
eaters probably fall between these two values (C. Pennycuick 1979). Two alternatives are
modeled. On pure scavenging trips, e = 11,480,000 joules/individual. In mixed fruitforaging (60%) and scavenging (33%) trips, e = 5,682,600 joules/individual at a minimum. Oldowan walking velocity is set at 1.1 m/sec (the low end of the optimally efficient
range for modern hominids: Margaria et al. 1963), since smaller hominids most efficient
velocity is lower than that of larger hominids. Basal metabolic rate (Em) is derived following C. Pennycuick (1979:168):
(2) Em = 4.8 mo.74
Em = 66.7 watts for a 35-kg hominid
where Em = basal metabolic rate in watts
m = body weight in kilograms
k is no larger than the empirically determined cost of walking at 1.1 m/sec for modern
humans twice as large as Oldowans (100.5 joules/kg/m: Taylor 1977; Margaria et al.
1963) and is probably smaller. k is derived (C. Pennycuick 1979:165):

r =

(3) k =


k = 91.4
where Elz = incremental cost of locomotion (cost of walking)
Substituting these values into equation (1) yields the values of r for a lone Oldowan
and for an Oldowan mother carrying a 10-kg nursing infant (Table 3). The latter circumstance is modeled as a worst-case scenario. Her metabolic rate is increased to 100.5 watts
by lactation (Crampton and Lloyd 1959; Portman 1970), but e, V, and k (Goldman and
Iampietro 1962) do not change. If an area of radius r were effectively searched annually,
by covering a 1 sector for each of 360 days (leaving 5 days/yr for repeat trips or illness),
then the foraging area and the total number of kilocarcasses encountered (kg-ctotal) can
be calculated (Table 4).
Whether or not kg-ctotal were sufficient for a scavenging Oldowan depends on other
species consumption, the Oldowans dietary needs, and the density of Oldowans within
the foraging area.
A crude estimate of kg-ceaten by nonhominids is derived from modern data (Houston
1979).Today, the mammalian predators eat only 35% of the yearly carcass biomass, with
vultures taking another So%, and the rest being either wasted or consumed by inverte-




Table 3
Energy extracted from a full gut load (e) and foraging radius (r) of Oldowans, at varying
rates of scavenging.

Lone Oldowan
100% scavenging
33% scavenging
Oldowan with infant
100% scavenging
33% scavenging

Value of c
(35 ka Oldowan)

Foraging radius,

Time spent daily

traveling. 2r

1 1,480,000joules

18.9 km
9.3 km

9 hr, 36 min
4 hr, 42 min

1 1,480,000joules

15.7 km
7.8 km

7 hr, 54 min
3 hr, 54 min

Table 4
Kilocarcassesencounteredby an Oldowan during foraging trips yearly and daily.

Lone Oldowan
100% meat
33% meat
Oldowan with infant
100% meat
33% meat

radius r

Foraging area
yearly (daily)


in km

in km'

yearly (daily)


1122 (3.1)
272 (0.76)

2,208,096 (16.8)
535,296 (4.1)


774 (2.2)
191 (0.53)

1,523,232 ( 1 1.6)
375,888 (2.9)

Table 5
Acquisition rates for a lone Oldowan on a pure scavenging trip (100%meat).

Foraging area

Maximum intake
(0.5 kg)
3.1 kmz

Minimum intake
(0.2 kg)
3.1 km'
7 Yo

brates. East's (1984) work shows that the relative proportions of herbivores and carnivores remain nearly constant with increased rainfall in savannahs like the Serengeti.
Thus, the percentage consumption by modern carnivores or their equivalents remained constant over the last 2 million years. Avian consumption is assumed to also scale with rainfall, accounting for about 30% of the dead herbivore biomass. Invertebrate consumption,
now at about 35%, varies with food availability; thus, that 35% is the maximum available for hominid scavenging.
Meat intake of spotted hyenas (0.66 kg of scavenged meat/individual/day: Kruuk
1972) and !Kung San (0.23 kg of meat/individual/day: Lee 1968) were used to set probable limits on intake by Oldowans. Because a lone Oldowan's metabolic rate was about
75% of these species' rates, these limits are scaled down to 0.5 and 0.2 kg meadindividual/day (Garland 1983). The Oldowan mother's intake was higher: 0.8 and 0.3 kg meat/
individuaVday. Meat intake by other Oldowans had only a small effect. Early hominid
densities are suggested to fall between 0.001 and 2 individuals/km2 when mixed diet (omnivory) is postulated (Boaz 1979; Martin 1981; Walker and Leakey 1978).
The daily acquisition rate (AR) needed by an Oldowan to maintain life is calculated




Table 6
Acquisition rates for a lone Oldowan on a mixed foraging-scavengingtrip (33%meat).

Foraging area

Maximum intake
(0.5 kg)
0.76 kmz

Minimum intake
(0.2 kg)
0.76 km2

7 yo

Table 7
Acquisitionrates for a lactating Oldowan for a pure scavenging trip (100O/0 meat).

Foraging area

Maximum intake
(Mother:0.8 kg)
(Others: 0.5 kg)
2.2 km2

Minimum intake
(Mother: 0.3 kg)
(Others: 0.2 kg)
2.2 km2

7 yo


3 yo

Table 8
Acquisition rates for a lactating Oldowan for a mixed foraging-scavengingtrip (33%meat).

Foraging area

Maximum intake
(Mother: 0.8 kg)
(Others: 0.5 kg)
0.53 km2

Minimum intake
(Mother: 0.3 kg)
(Others: 0.2 kg)
0.53 km2
1 1 OO/

as a percentage of kg-ctotal encountered within its foraging radius; ARgroup is the total
needed by Oldowans living at a maximum density and sharing a foraging radius (Tables

A lone Oldowan needed to obtain 1%-12% of the kilocarcasses available for scavenging in its daily foraging area if no other hominids were present or 7%-9% if others shared
the area. In these cases, acquisition rates were much lower than 35%, indicating that
scavenging by Oldowans was feasible. Further, the acquisition rates are so low that there
is a wide margin for error in the estimates and assumptions without rendering scavenging
unfeasible for an Oldowan (contra Schaller and Lowther 1969).
The case most closely approaching the limits of the system is that of an Oldowan
mother with nursing infant (Tables 7-8). If she had the maximum postulated meat intake, and if she lived in an area of high hominid density, she had an AR of 29%. Given
that all estimates were made to test the feasibility of the scavenging hypothesis severely,
while remaining within the bounds of reason, it is remarkable that the AR remains below
35% even in this most difficult situation. This reconstruction does not require either cooperative social behavior or provisioning of females and offspring. It is concluded that
the predictions of feasibility are met.




Conclusions: Scavenging, the Home Base Hypothesis and Human Evolution

All tests of predictions of the scavenging hypothesis given here are fulfilled by a generous margin. It is concluded that the scavenging hypothesis is not refuted and is worthy
of additional investigation.
The idea that scavenging may have been a major food-procurement strategy in Bed I
times has broader implications. The first of these is that it casts doubt on the idea that
sites represent central places, home bases, or camp sites that were the focus of transport
and sharing of food. The cut-mark evidence strongly suggests that systematic disarticulation to enable transport and sharing of carcasses did not occur. The feasibility data
suggest that provisioning, food sharing, and division of labor are not necessarily intrinsic
to strategies involving utilization of carcasses. Additional problems with the familiarly
human interpretation of Bed I sites arise because of the many biological and ecological
differences between Oldowans and modern hunter-gatherers. Since Oldowans were small
and relatively small-brained, lacked fire, projectile weapons, and domestic dogs, and
lived in areas with a higher large carnivore density than any modern African habitat, the
danger of sleeping and caring for offspring near carcasses was considerably greater for
Oldowans than for modern humans (Shipman 1983). In short, it is difficult to reconcile
this information with the notion that Bed I sites were base camps.
Other evidence leads to questions about the home base hypothesis, too. Potts (1982,
1984) presents other evidence that competition from carnivores seriously restricted 01dowan access to carcasses and made the time spent in carcass-processing areas likely to
be brief. He reports that the disparate weathering on Bed I bones indicates either occupation over four or more years at each site or repeated reoccupation; neither of these
behaviors is common among modern hunter-gatherers.
If Bed I sites are not base camps, what mechanism accounts for the abnormally dense
accumulations of bones associated with stone tools? Two possibilities that are not mutually exclusive are suggested. The first is that Bed I sites represent safe areas, such as
trees, to which scavenged items were carried. Bones and tools accumulated as these safe
areas were used repeatedly. Unlike hunting, transport of scavenged items does not invariably require disarticulation with tools, since body parts or scraps separated from the
carcass by primary predators are prime targets for scavenging. The second possibility,
articulated by Potts (1982, 1984), is that these sites represent stone caches, to which resources were taken for processing. Although Potts assumes random distributions of both
caches and faunal resources, and I make a similar assumption about carcasses, neither
carcasses nor faunal resources are randomly distributed (Blumenschine 1984; Behrensmeyer and Dechant-Boaz 1980). The discrepancy between our models and reality means
that Oldowans could have improved the frequency with which they encountered carcasses/faunal resources by utilizing their habitat selectively. The stone cache hypothesis
is especially congruent with the scavenging hypothesis if caches were placed or used more
frequently near features that favored scavenging. Such features might include trees suitable for climbing, or waterholes, because of the high frequency ofcarcasses and the habits
of carnivores (Binford 1984).
A major site from a later time period has been interpreted as preserving scavenged
remains: Klasies River Mouth (KRM), reinterpreted by Binford (1984) following upon
quite different interpretations by Klein (1981 and references therein). Whichever interpretation is correct, the KRM data differ in interesting ways from the Bed I data. First,
at KRM there was differential treatment of bovids according to size class. Binford maintains that larger bovids (size classes IV-V) were scavenged, whereas smaller ones (size
classes 1-111) were hunted. This pattern is not apparent at Olduvai: the sample of cutmarked, apparently scavenged bones includes only one larger bovid specimen. Second,
the KRM scavenged material was apparently processed heavily for marrow, whereas the
Olduvai bones were not intensively utilized and modified for marrow extraction. Potts
(1984:344) concurs, stating that Olduvai bones show incomplete processing . . . [and]



[88, 1986

suggest that hominids abandoned considerable portions of meat and marrow at each
site. Finally, Binford (1984~86)remarks on the inflexible behavior of KRM hominids
in scavenging: an almost stimulus-response structure of behavior. My impression is
that the Olduvai data do not reveal strong patterning or routinized treatment of remains,
but rather more opportunistic and haphazard processing. These differences may indicate
either that scavenging behaviors changed as hominids evolved or that the interpretation
of one of the two faunas (KRM or Olduvai) as scavenged is inaccurate.
Finally, why bipedalism arose is a classic issue. The striking congruency between the
attributes of bipedalism, as analyzed here, and the locomotor needs of scavengers might
suggest to some that bipedalism is actually an adaptation, not an exaptation (Gould and
Vrba 1982), to scavenging. If it is to be concluded that the origins of bipedalism and
scavenging are causally related, one of two difficulties must be surmounted. Either the
earliest evidence for toolmaking must be pushed back to more closely approximate that
of bipedalism or it must be postulated that effective scavenging was possible without dental or technological adaptations for carcass processing.
Acknowledgments. This work was funded by the National Science Foundation (BNS 80- 1397 and
80-2-1397) and from the Boise Fund. I appreciate the assistance and cooperation of the Presidents
Office of Kenya (permit OP. 13/001/6C70), the Government ofTanzania and the staff of the National Museums of Kenya. Especial thanks go to M. D., R. E., and M. E. Leakey for their help.
This paper benefited greatly from the advice and discussion ofJ. Buikstra, M. Cartmill, G. Isaac,
R. Lewin, R. Potts, M. Schoeninger, R. Smith, M. Teaford, R. Taylor, B. Van Valkenburgh, A.
Walker, and J. Rose. J. Rose and B. Coe provided technical assistance.

References Cited
Ardrey, R.
1961 African Genesis: A Personal Investigation into the Animal Origins and Nature of Man.
New York: Dell.
1976 The Hunting Hypothesis: A Personal Conclusion Concerning the Evolutionary Nature of
Man. New York: Atheneum Press.
Behrensmeyer, A. K., and D. E. Dechant-Boaz
1980 The Recent Bones ofAmboseli Park, Kenya, in Relation to East African Paleoecology. In
Fossils in the Making. A. K. Behrensmeyer and A. P. Hill, eds. Pp. 72-93. Chicago: University
of Chicago Press.
Bertram, B. C. R.
1973 Lion Population Regulation. East African Wildlife Journal 1 1:215-225.
1975 Weights and Measures of Lions. East African Wildlife Journal 13:141-143.
1979 Serengeti Predators and Their Social Systems. In Serengeti: Dynamics of an Ecosystem.
A. R. E. Sinclair and M. Norton-Griffiths, eds. Pp. 221-248. Chicago: University of Chicago
Bicchieri, M. D., ed.
1972 Hunters and Gatherers Today. New York: Holt, Rinehart & Winston.
Binford, L. R.
1981 Bones: Ancient Men and Modern Myths. San Francisco: Academic Press.
1984 Faunal Remains from Klasies River Mouth. San Francisco: Academic Press.
Blumenschine, R. J.
1984 Scavenging in the Serengeti. Anthroquest 28:ll-12.
Boaz, N. T.
1979 Early Hominid Population Densities: New Estimates. Science 206:592-595.
Bonnefille, R.
1977 Etudes palynologiques des depots Plio-Pleistocene (Beds I and 11). National Geographic
Society Report.
Brain, C. K.
1970 New Finds at the Swartkrans Australopithecine Site. Nature 225:1112-1119.
1981 The Hunters or the Hunted? An Introduction to African Cave Taphonomy. Chicago:
University of Chicago Press.




Bunn, H.
1982 Meat-eating and Human Evolution. Ph.D. thesis, University of California, Berkeley.
1983a Comparative Analysis of Modern Bone Assemblages from a San Hunter-Gatherer
Camp in the Kalahari Desert, Botswana, and from a Spotted Hyena Den near Nairobi, Kenya.
In Animals and Archaeology, Vol. 1. J. Clutton-Brock and C. Grigson, eds. Pp. 143-148. London: British Archaeological Reports.
1983b Evidence on the Diet and Subsistence Patterns of Plio-Pleistocene Hominids at Koobi
Fora, Kenya, and at Olduvai Gorge, Tanzania. In Animals and Archaeology, Vol. 1. J. Clutton-Brock and C. Grigson, eds. Pp. 2 1-30. London: British Archaeological Reports.
Ceding, T., R. Hay, and J. ONeil
1977 Isotopic Evidence for Dramatic Climatic Changes in East Africa during the Pleistocene.
Nature 267:137-138.
Coe, M. J., D. H. Cumming, and J. Phillipson
1976 Biomass and Production of Large African Herbivores in Relation to Rainfall and Primary
Production. Oecologia 22:34 1-354.
Coon, C. S.
1971 The Hunting Peoples. Harmondsworth, England: Pelican Books.
Crader, D. C.
1983 Recent Single-carcass Bone Scatters and the Problems of Butchery Sites in the Archaeological Record. In Animals and Archaeology, Vol. 1. J. Clutton-Brock and C. Grigson, eds.
Pp. 107-141. London: British Archaeological Reports.
Crampton, E., and L. Lloyd
1959 Fundamentals of Nutrition. San Francisco: W. H. Freeman.
Cronin, J. E., N. T. Boaz, C. S. Stringer, and Y. Rak
1981 Tempo and Mode of Hominid Evolution. Nature 292:113-122.
Curio, E.
1976 The Ethology of Predation. Berlin: Spring-Verlag.
Darwin, C.
1871 Descent of Man. London: John Murray & Sons.
Dart, R. A.
1959 Adventures with the Missing Link. Philadelphia: Institutes Press.
Dunbar, R. I. M.
1983 Theropithecines and Hominids: Contrasting Solutions to the Same Ecological Problem.
Journal of Human Evolution 12:647-658.
East, R.
1984 Rainfall, Soil Nutrient Status and Biomass of Large African Savannah Animals. African
Journal of Ecology 22(4):245-270.
Eaton, R. L.
1979 Interference Competition among Carnivores: A Model for the Evolution of Social Behavior. Carnivore 29-16.
Estes, R., and J. Goddard
1967 Prey Selection and Hunting Behavior of the African Wild Dog. Journal of Wildlife Management 31( 1):52-70.
Ewer, R. F.
1973 The Carnivores. I thaca: Cornell University Press.
Fedak, M. A,, and H. J. Seeherman
1979 Re-appraisal of Energetics of Locomotion Shows Identical Cost in Bipeds and Quadrupeds including Ostrich and Horse. Nature 282:713-716.
Frame, L. H., and G. W. Frame
1976 Wild Dogs of the Serengeti. African Wildlife Foundation Newsletter (Nairobi) 11(3):1-6.
Garland, T.
1983 Scaling the Ecological Cost of Transport to Body Mass in Terrestrial Mammals. American Naturalist 121:571-587.
Gentry, A. W., and A. Gentry
1978a Fossil Bovidae (Mammalia) of Olduvai Gorge, Tanzania, Part I. Bulletin of the British
Museum of Natural History (Geology) 2 9 ( 4 ) : 2 8 W .
1978b Fossil Bovidae (Mammalia) ofOlduvai Gorge, Tanzania, Part 11. Bulletin of the British
Museum of Natural History (Geology) 30( 1): 1-83.



[88, 1986

Gifford, D.
1977 Observations of Modern Human Settlements as an Aid to Archaeological Interpretation.
Ph.D. dissertation, University of California, Berkeley.
Gifford, D. P., G. L1. Isaac, and C. M. Nelson
1981 Evidence for Predation and Pastoralism at Prolonged Drift: A Pastoral Neolithic Site in
Kenya. Azania 15:57-108.
Goldman, R. F., and P. F. Iampietro
1962 Energy Cost of Load Carriage. Journal of Applied Physiology 17:675-676.
Could, R. A.
1968 Living Archaeology: The Ngatatjara of Western Australia. Southwest Journal of Anthropology 24: 104-122.
1980 Living Archaeology. Cambridge: Cambridge University Press.
Could, S.J., and E. S. Vrba
1982 Exaptation: A Missing Term in the Science of Form. Paleobiology 8:4-15.
Guilday, J. E., P. W. Parmalee, and D. P. Tanner
1962 Aboriginal Butchering Techniques at the Eschelman Site (36LA12), Lancaster County,
Pennsylvania. Pennsylvania Archaeology 32:59-83.
Hay, R.
1976 Geology of the Olduvai Gorge: A Study of Sedimentation in a Semiarid Basin. Berkeley:
University of California Press.
Hewes, G. W.
1961 Food Transport and the Origin of Hominid Bipedalism. American Anthropologist
63:687-7 10.
1964 Hominid Bipedalism: Independent Evidence for the Food-carrying Theory. Science
Hill, K.
1982 Hunting- and Human Evolution..lournal
of Human Evolution 11:521-544.
Holloway, R. L.
1978 Problems of Brain Interpretation and African Hominid Evolution. In Early Hominids of
Africa. C. J.Jolly, ed. Pp. 379401. New York: St. Martins Press.
Houston, D.C.
1979 The Adaptations of Scavengers. In Serengeti: Dynamics of an Ecosystem. A. R. E. Sinclair
and M. Norton-Griffiths, eds. Pp. 263-286. Chicago: University of Chicago Press.
Howells, W.
1959 Mankind in the Making: The Story of Human Evolution. Garden City, NY: Doubleday.
Isaac, G. L1.
1978 The Food-sharing Behavior of Proto-human Hominids. Scientific American 238:9&108.
1983 Bones in Contention: Competing Explanations for the Juxtaposition of Early Pleistocene
Artifacts and Faunal Remains. In Animals and Archaeology, Vol. 1. J. Clutton-Brock and C.
Grigson, eds. Pp. 3-19. London: British Archaeological Reports.
Isaac, G. L l . , and D. Crader
1981 To What Extent Were Early Hominids Carnivorous? An Archaeological Perspective. In
Omnivorous Primates: Gathering and Hunting in Human Evolution. R. S. 0. Harding and
G. Teleki, eds. Pp. 37-103. New York: Columbia University Press.
Johanson, D. C., and T. D. White
1979 A Systematic Assessment of Early African Hominids. Science 203:321-330.
Johanson, D. C., M. Taieb, and Y. Coppens
1982 Pliocene Hominids from the Hadar Formation, Ethiopia (1973-1977): Stratigraphic,
Chronologic and Paleoenvironmental Contexts, with Notes on Hominid Morphology and Systematics. American Journal of Physical Anthropology 57:373402.
Jolly, C. J.
1970 The Seed-eater Hypothesis: A New Model ofHominid Differentiation Based on a Baboon
Analogy. Man 55-20.
Klein, R. G.
1981 Stone Age Predation of Small African Bovids. South African Archaeological Bulletin
Kruuk, H.
1972 The Spotted Hyena: A Study of Predation and Social Behavior. Chicago: University of
Chicago Press.
1976 Feeding and Social Behavior of the Striped Hyaena (Hyaenavulgaris Desmarest). East African WildlifeJournal 14(2):91-112.





Leakey, M.D.
1971 Olduvai Gorge, Vol. 3. London: Cambridge University Press.
Leakey, M. D., and R. L. Hay
1979 Pliocene Footprints in the Laetolil Beds at Laetoli, Northern Tanzania. Nature 278:317323.

Lee, R. B.
1968 What Hunters Do for a Living, or, How to Make Out on Scarce Resources. In Man the
Hunter. R. E. Lee and I. De Vore, eds. Pp. 30-48. Chicago: Aldine.
Lee, R. B., and I. DeVore, eds.
1968 Man the Hunter. Chicago: Aldine.
Lovejoy, C. 0.
1974 The Gait ofAustralopithecines. Yearbook of Physical Anthropology 41:191-215.
Lovejoy, C. O., K. G. Heiple, and A. H. Burstein
1973 The Gait of Australopithccus. American Journal of Physical Anthropology 38:757-780.
Margaria, R., P. Cerretelli, P. Aghemo, and G. Sassi
1963 Energy Cost of Running. Journal of Applied Physiology 18:367-370.
Marks, S. A.
1976 Large Mammals and a Brave People. Seattle: University of Washington Press.
Marshall, L.
1965 The !Kung Bushmen of the Kalahari Desert. In Peoples of Africa. J. L. Gibbs, ed. Pp.
241-278. New York: Holt, Rinehart & Winston.
Martin, R. A.
1981 On Extinct Population Densities. Journal of Human Evolution 10:427-428.
McHenry, H. M.
1976 Early Hominid Body Weight and Encephalization. American Journal of Physical Anthropology 45:77-84.
1978 Fore- and Hindlimb Proportions in Plio-Pleistocene Hominids. American Journal of
Physical Anthropology 4 3 : 3 9 4 .
1982 The Pattern of Human Evolution: Studies on Bipedalism, Mastication, and Encephalization. Annual Review of Anthropology 11:151-173.
McHenry, H. M., and L. A. Temerin
1979 The Evolution of Hominid Bipedalism: Evidence from the Fossil Record. Yearbook of
Physical Anthropology 22:105-131.
Norton-Griffiths, M., D. Herlocker, and L. Pennycuick
1975 The Patterns of Rainfall in the Serengeti Ecosystem, Tanzania. East African Wildlife
Journal 13:347-374.
Nowak, R. M., and J. L. Paradiso
1983 Walkers Mammals of the World, 4th ed. Baltimore: Johns Hopkins University Press.
Oliver, J. S.
1985 Bone Damage Morphologies from Shield Trap Cave, Carbon County, Montana. Abstracts, First International Conference on Bone Modification, p. 27.
Pennycuick, C. J.
1971 The Soaring Flight of Vultures. Scientific American 229: 102-109.
1979 Energy Costs of Locomotion and the Concept of Foraging Radius. In Serengeti: Dynamics of an Ecosystem. A. R. E. Sinclair and M. Norton-Grifiths, eds. Pp. 164-184. Chicago:
University of Chicago Press.
Pennycuick, L.
1975 Movements of the Migratory Wildebeest Population in the Serengeti Area between 1960
and 1973. East African Wildlife Journal 13(1):65-87.
Portman, 0.
1970 Nutritional Requirements (NCR) of Non-Human Primates. In Feeding and Nutrition of
Nonhuman Primates. R. Harris, ed. Pp. 87-1 16. New York: Academic Press.
Potts, R.
1982 Lower Pleistocene Site Formation and Hominid Activities at Olduvai Gorge, Tanzania.
Ph.D. dissertation, Harvard University, Cambridge.
1984 Home Bases and Early Hominids. American Scientist 72:336347.
Potts, R., and P. Shipman
1981 Cutmarks Made by Stone Tools on Bones from Olduvai Gorge, Tanzania. Nature




Prost, J. H.
1980 Origin of Bipedalism. American Journal of Physical Anthropology 52: 175-189.
Robinson, J. T.
1972 Early Hominid Posture and Locomotion. Chicago: University of Chicago Press.
Rodman, P., and H . M. McHenry
1980 Bioenergetics and the Origin of Hominid Bipedalism. American Journal of Physical Anthropology 52: 103-106.
Rose, J. J.
1983 A Replication Technique for Scanning Electron Microscopy: Applications for Anthropologists. American Journal of Physical Anthropology 62:255-263.
Rose, M. D.
1984 Food Acquisition and the Evolution of Positional Behavior: The Case of Bipedalism. In
Food Acquisition and Processing in Primates. D. T. Chivers, B. A. Wood, and A. Bilsborough,
eds. Pp. 509-524. New York: Plenum Press.
Rudnai, J.
1973 The Social Life of the Lion. Wallingford, PA: Washington Square East, Publishers.
Savage, R. J. G.
1978 Carnivora. I n Evolution of African Mammals. V. J. Maglio and H. B. S. Cooke, eds. Pp.
249-267. Cambridge, MA: Harvard University Press.
Schaller, G.
1968 Hunting Behavior of the Cheetah in the Serengeti National Park, Tanzania. East African
WildlifeJournal 6:95-100.
1972 The Serengeti Lion: A Study of Predatory-Prey Relations. Chicago: University ofChicago
Schaller, G., and G. Lowther
1969 The Relevance of Carnivore Behavior to the Study of Early Hominids. Southwest Journal
of Anthropology 25(4):307-341.
Senut, B.
1981 Humeral Outlines in Some Hominoid Primates and in Plio-Pleistocene Hominids. American Journal of Physical Anthropology 56:275-283.
Service, E. R.
1979 The Hunters. Englewood Cliffs, N.J.: Prentice-Hall.
Shipman, P.
1981 Applications of Scanning Electron Microscopy to Taphonomic Problems. In The Research Potential of Anthropological Museum Collections. Annals of the New York Academy
of Sciences 276:357-385.
1983 Early Hominid Lifestyle: Hunting and Gathering or Foraging and Scavenging? In Animals and Archaeology, Vol. 1. J. Clutton-Brock and C. Grigson, eds. Pp. 31-50. London: British Archaeological Reports.
1984 Scavenger Hunt. Natural History 93(4):2&27.
Shipman, P., and J. J. Rose
1983a Early Hominid Hunting, Butchering and Carcass-processing Behaviors: Approaches to
the Fossil Record. Journal of Anthropological Archaeology 2( 1):57-98.
1983b Evidence of Butchery and Hominid Activity at Torralba and Ambrona: An Evaluation
Using Microscopic Techniques. Journal of Archaeological Science 10(3):465-474.
Sinclair, A. R.E.
1979 The Serengeti Environment. In Serengeti: Dynamics of an Ecosystem. A. R. E. Sinclair
and M.Norton-Griffiths, eds. Pp. 31-45. Chicago: University of Chicago Press.
Stern, J. T., and R. L. Susman
1983 The Locomotor Anatomy of Australopithccus afarensis. American Journal of Physical Anthropology 60:279-317.
Steudel, K.
1980 New Estimates of Early Hominid Body Size. American Journal of Physical Anthropology

Susman, R. L., and J. T. Stern

1979 Telemetered Electromyography of Flexor Digitorum Profundus and Flexor Digitorum
Superficialis in Pan troglo&es and Implications for Interpretation of the 0.H. 7 Hand. American Journal of Physical Anthropology 50:565-574.
1982 Functional Morphology of Homo habilis. Science 2 17:931-934.




Szalay, F.
1975 Hunting-Scavenging Protohominids: A Model for Hominid Origins. Man 10:420-429.
Taylor, C. R.
1977 The Energetics of Terrestrial Locomotion and Body Size in Vertebrates. In Scale Effects
in Animal Locomotion. T. J. Pedley, ed. Pp. 127-141. London: Academic Press.
Taylor, C. R., N. C. Heglund, and G. M. 0. Maloiy
1982 Energetics and Mechanics of Terrestrial Locomotion, 1: Metabolic Energy Consumption
as a Function of Speed and Body Size in Birds and Mammals. Journal of Experimental Biology
Tiger, L., and R. Fox
1971 The Imperial Animal. New York: Holt, Rinehart & Winston.
Turnbull, C.
1965a The Mbuti Pygmies of the Congo. In Peoples of Africa. J. L. Gibbs, ed. Pp. 279-318.
New York: Holt, Rinehart & Winston.
1965b Wayward Servants: The Two Worlds of the African Pygmies. Garden City, NY: Natural
History Press.
Van Lawick-Goodall, J., and H. Van Lawick
1970 Innocent Killers. London: Collins.
Vrba, E. S.
1979 A New Study of the Scapula ofAustralopithccusahcanus from Sterkfontein. American Journal of Physical Anthropology 5 1 : 1 17-1 29.
1980 The Significance of Bovid Remains as Indicators of Environment and Predation Patterns.
I n Fossils in the Making. A. K. Behrensmeyer and A. P. Hill, eds. Pp. 247-272. Chicago: University of Chicago Press.
Walker, A.
1980 Functional Anatomy and Taphonomy. In Fossils in the Making. A. K. Behrensmeyer and
A. P. Hill, eds. Pp. 182-196. Chicago: University of Chicago Press.
1981 Dietary Hypotheses and Human Evolution. Philosophical Transactions of the Royal Society of London (series B) 292:57-64.
1984 Extinction in Human Evolution. In Extinctions. M. Niteki, ed. Pp. 119-152. Chicago:
University of Chicago Press.
Walker, A., and R. E. F. Leakey
1978 The Hominids of East Turkana. Scientific American 239:54-66.
Washburn, S. L.
1960 Tools and Human Evolution. Scientific American 203:63-75.
1978 The Evolution of Man. Scientific American 239:194-208.
Washburn, S. L., and C. S. Lancaster
1968 The Evolution of Hunting. In Man the Hunter. R. B. Lee and I. DeVore, eds. Pp. 293303. Chicago: Aldine.
Winterhalder, B., and E. A. Smith, eds.
1981 Hunter-gatherer Foraging Strategies: Ethnographic and Archaeological Analyses. Chicago: University of Chicago Press.
Yellen,J. E.
1977 Cultural Patterning in Faunal Remains: Evidence from the !Kung Bushmen. I n Experimental Archaeology. D. Ingersoll, J. E. Yellen, and W. Macdonald, eds. Pp. 271-331. New
York: Columbia University Press.
Zihlman, A. L.
1978 Interpretations of Early Hominid Locomotion. In Early Hominids of Africa. C. J. Jolly,
ed. Pp. 361-377. London: Duckworth.
Zihlman, A. L., and L. Brunker
1979 Hominid Bipedalism: Then and Now. Yearbook of Physical Anthropology 22: 132-162.