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PATSHIPMAN

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.
ARLY HOMINID LIFE IS OFTEN RECONSTRUCTED AS BROADLY SIMILAR

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
PATSHIPMAN
is Assistant Prgessor,Department of Cell Biology and Anatomy, School of Mcdicinc,Johns Hopkins University,
Baltimore, M D 21205.

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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

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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
agent.

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
level.
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;

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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
criticism.
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
1983a).
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
behavior.
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

SCAVENGING OR HUNTING
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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
26

Midshaft
34

272

24

28

42

(b) Prolonged Drift


(c) Olduvai tooth marks

Significance
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
Significance
22
38
avs. b: chiz = 2.5
(a) Olduvai cut marks
1 degree of freedom
p > 0.05
(b) Prolonged Drift
77
217
b vs. c: chi2 = 50.9
1 degree of freedom
p C 0.005*
13
a vs. c: chiz = 17.9
(c) Olduvai tooth marks
41
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

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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
1973).
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.

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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
point.
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

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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.

eV
4 Em 4kV
e
=
the energy extracted from a full gut load
where
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):
(1)

r =

(3) k =

V
El2

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-

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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,
T

Time spent daily


traveling. 2r

1 1,480,000joules
5,682,600joules

18.9 km
9.3 km

9 hr, 36 min
4 hr, 42 min

1 1,480,000joules
5,682,600joules

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

Foraging
radius r

Foraging area
yearly (daily)

Kilocarcasses

in km

in km'

yearly (daily)

18.9
9.3

1122 (3.1)
272 (0.76)

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

15.7
7.8

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
Kg-ctotal
AR
ARgroup

Maximum intake
(0.5 kg)
3.1 kmz
16.8
3'/o
18%

Minimum intake
(0.2 kg)
3.1 km'
16.8
1Yo
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

36

[88,1986

AMERICAN
ANTHROPOLOGIST

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

Foraging area
Kg-ctotal
AR
ARgroup

Maximum intake
(0.5 kg)
0.76 kmz
4.1
12%
19%

Minimum intake
(0.2 kg)
0.76 km2
4.1

5%
7 yo

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

Foraging area
Kg-ctotal
AR
ARgroup

Maximum intake
(Mother:0.8 kg)
(Others: 0.5 kg)
2.2 km2
11.6

Minimum intake
(Mother: 0.3 kg)
(Others: 0.2 kg)
2.2 km2
11.6

7 yo
22%

8%

3 yo

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

Foraging area
Kg-ctotal
AR
ARgroup

Maximum intake
(Mother: 0.8 kg)
(Others: 0.5 kg)
0.53 km2
2.9
28%
29%

Minimum intake
(Mother: 0.3 kg)
(Others: 0.2 kg)
0.53 km2
2.9
10%
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

5-8)*
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.

Shipman]

SCAVENGING OR HUNTING
IN EARLYHOMINIDS

37

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]

38

AMERICAN
ANTHROPOLOGIST

[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.

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