Professional Documents
Culture Documents
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
27
28
AMERICAN
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[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
Shipman]
SCAVENGING OR HUNTING
IN EARLYHOMINIDS
29
30
AMERICAN
ANTHROPOLOGIST
[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
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.
SCAVENGING OR HUNTING
IN EARLYHOMINIDS
Shipman]
31
Table 1
Distribution of cut marks and tooth marks on Olduvai bovids relative to joints compared
with that of cut marks from Prolonged Drift.
Near joint
26
Midshaft
34
272
24
28
42
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
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*
~
~~
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.
32
AMERICAN
ANTHROPOLOGIST
[88,1986
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.
Shipman]
SCAVENGING OR HUNTING
IN EARLYHOMINIDS
33
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.
34
AMERICAN
ANTHROPOLOGIST
[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.
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-
Shipman]
SCAVENGING OR HUNTING
IN EARLYHOMINIDS
35
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
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
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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