Journal of Archaeological Science 34 (2007) 2025e2034

http://www.elsevier.com/locate/jas

Carnivore competition, bone destruction, and bone density

J. Tyler Faith a,*, Curtis W. Marean b, Anna K. Behrensmeyer c
a
Hominid Paleobiology Doctoral Program, CASHP, Department of Anthropology, George Washington University, 2110 G St NW,
Washington, DC 20052, USA
b
Institute of Human Origins, School of Human Evolution and Social Change, PO Box 872402, Arizona State University,
Tempe, AZ 85287, USA
c
Department of Paleobiology, Evolution of Terrestrial Ecosystems Program, Smithsonian Institution, National Museum of Natural History,
PO Box 37012, NHB, MRC-121, Washington, DC 20013-7012, USA
Received 25 September 2006; received in revised form 24 December 2006; accepted 2 January 2007

Abstract

In carnivore-modified archaeofaunal assemblages it is important to evaluate the degree to which carnivores have overprinted hominin behav-
ioral signals. To examine the signals of increased competition for discarded bone, we present controlled experimental data on 33 simulated ar-
chaeological sites subjected to secondary consumption by spotted hyenas. We examine the relationship between competition, as measured by
controlled numbers of hyenas and limb bones, and resultant levels of destruction and correlations between long-bone portion survivorship and
bone density. Our results indicate that levels of destruction are equivalent regardless of the numbers of hyenas or long-bones included in the
experimental assemblages. Correlations between long-bone epiphyseal and near-epiphyseal portions and bone density, however, do provide
an indication of the level of competition. Results from the experimental study are used to highlight divergent levels of carnivore competition
for hominin-discarded bone at the Plio-Pleistocene localities FLKN-Zinjanthropus and FLKN levels 1e2 from Olduvai Gorge, Tanzania.
Ó 2007 Elsevier Ltd. All rights reserved.

Keywords: Carnivore competition; Carnivore destruction; Bone density; Spotted hyena; Taphonomy; Olduvai

1. Introduction characteristics - primarily bone density (Brain, 1967, 1969;

Reconstructing the taphonomic history of an archaeofaunal
assemblage requires an assessment of the processes that de-
stroy and alter faunal remains following human discard. An
evaluation of these processes, which includes, but is not lim-
ited to, subaerial weathering (Behrensmeyer, 1978), carnivore
modification (Binford et al., 1988; Blumenschine, 1988;
Blumenschine and Marean, 1993; Marean and Spencer,
1991; Marean et al., 1992), and post-depositional alteration
(Klein and Cruz-Uribe, 1984; Marean, 1991), is a crucial
step toward inferring hominin behavioral patterns. It has
long been recognized that differential survivorship of skeletal
elements and portions thereof is mediated by physical
* Corresponding author. Tel.: þ1 202 994 0154.
E-mail address: tfaith@gwu.edu (J.T. Faith).
Lam et al., 1998; Lyman, 1994). Since Lyman’s (1984)
pioneering study of bone density in deer (Odocoileus spp.),
bone density analyses of archaeofaunal assemblages have
provided a means of gauging the effects of density-mediated
attrition on skeletal element representation. The presence of
a significant positive correlation between skeletal element
abundance and bone density, when an assemblage has not
been subjected to biasing collection procedures, is presumed
to reflect density-mediated destruction of faunal remains
(Grayson, 1989; Klein, 1989; Lam et al., 1998; Lupo, 1995;
Lyman, 1993, 1994).
Of the many destructive processes considered to be affected
by bone density, carnivore destruction has been documented in
experimental contexts to be at least partially density-mediated
(Marean and Spencer, 1991; Marean et al., 1992). Carnivore
destruction can include both primary destruction of bones en-
countered from carcasses preyed upon by carnivores, and

0305-4403/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2007.01.017
2026 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034
secondary consumption of skeletal portions previously dis-
carded by human foragers. Low-density elements and portions
of bone, such as long-bone epiphyses, retain an abundance of
bone grease distributed within cancellous bone that is highly at-
tractive to carnivores (Binford, 1978; Blumenschine, 1988;
Lyman, 1985). The abundances of skeletal elements in faunal
assemblages subjected to carnivore destruction are thus ex-
pected to correlate positively with bone density. This is not
always the case (Carlson and Pickering, 2004; Pickering and
Carlson, 2002), and it has been suggested that the absence of
a correlation in situations of undoubted carnivore interaction
with a bone assemblage may relate to the use of inaccurate
bone density data or a violation of the conditions required for
a correlation analysis (Lam and Pearson, 2005). Alternatively,
a weak correlation between bone survivorship and bone density
may reflect local ecological factors such as the availability of
consumable bone and intensity of carnivore competition (Faith
and Behrensmeyer, 2006). Here we present new information on
carnivore destruction of bone in an examination of experimen-
tal patterns of bone destruction as they relate to bone density
and varying levels of conspecific competition.
Previous experimental studies have subjected ‘‘simulated
archaeological sites’’ (as in Blumenschine, 1988) to spotted
hyena modification to show that middle shaft portions of
long-bones provide the most accurate estimate of the original
number of limb elements (Marean and Spencer, 1991) and to
present controlled data on the sequence of bone choice and dif-
ferential levels of destruction to axial, appendicular, and com-
pact bones (Marean et al., 1992). In this analysis we expand
on these studies and use Marean’s original experimental data
to examine the relationship between levels of carnivore compe-
tition and the differential survivorship of appendicular bone
portions of varying densities. This allows us to examine skeletal
element abundance and bone density correlations under con-
trolled levels of competition, as measured by a predetermined
number of hyenas and limb elements.
In a previous paper, Faith and Behrensmeyer (2006) sug-
gested that the strength of the correlation between long-bone
epiphyses survival and bone density reflects the intensity of car-
nivore competition and degree of modification to an assem-
blage. This proposal is based on an actualistic case study of
patterns of skeletal element survival under changing abundances
of spotted hyena (Crocuta crocuta), an effective bone-cruncher,
in Amboseli Park, Kenya. At times with low abundances of Cro-
cuta, long-bone epiphysis survival correlates significantly with
bone density. When the Crocuta population in Amboseli
increased seven-fold, the correlation declines to insignificance.
Faith and Behrensmeyer interpret this pattern as a function of
a carnivore’s ability to be selective in choosing low-density ele-
ments and portions for consumption. At low levels of competi-
tion (e.g., an individual carnivore consuming carcass remains),
carnivores can be selective. When inter- or intra-specific compe-
tition is high (e.g., a feeding frenzy), carnivores do not discrim-
inate between high-and low-density portions, resulting in a poor
correlation with bone density. In such situations, little survives
other than the highest density pieces completely lacking cancel-
lous bone.
Carnivores have been implicated in the taphonomic histories
of archaeological bone assemblages in a broad range of
geographic and environmental contexts (e.g., Assefa, 2006;
Bartram and Marean, 1999; Blumenschine, 1995; Brain,
1981; Chase et al., 1994; Domı́nguez-Rodrigo et al., 2002;
Marean et al., 2000; Marean and Kim, 1998; Monahan, 1996;
Mondini, 2002; Potts, 1988). Evaluating the levels of carnivore
competition for human-discarded bone can be a powerful tool
for assessing the degree to which carnivores have overprinted
hominin behavioral signals. Although the patterns in selected
archaeological contexts closely match the patterns expected
based on the Amboseli data (Faith and Behrensmeyer, 2006),
this previous study lacks a direct observational link between
the processes (e.g., level of carnivore competition) and result-
ing patterns (e.g., strength of correlation between limb-end
abundance and bone density). A cross-validation of the rela-
tionship between carnivore competition and bone density in
controlled experimental settings strengthens the application
of the actualistic data from Amboseli to the archaeological
record (Marean, 1995). With this in mind, we present new anal-
yses of Marean’s experimental data (Marean and Spencer,
1991; Marean et al., 1992) to enhance our understanding of
the link between carnivore competition, bone destruction,
and bone density.
2. Experimental methods
The processes involved in carnivore modification of a bone
assemblage, termed ‘‘carnivore ravaging’’ by Binford (1981),
include bone choice and transportation, modification by chew-
ing, and destruction by consumption. Bunn (1986) and Bunn
and Kroll (1986) had observed that secondary carnivore con-
sumption of human-discarded bone resulted in deletion of
long-bone epiphyses. Marean and Spencer (1991) provided ex-
perimental and quantitative documentation that epiphyses and
near-epiphyseal fragments are consistently consumed and de-
stroyed by spotted hyenas in order to digest nutritious bone
grease within these cancellous portions. Here we test whether
variation in the destruction of long-bone portions and the rela-
tionship between surviving portions and bone density is af-
fected by competition for nutrients among different numbers
of bone consumers.
Simulated archaeological sites were designed so as to accu-
rately replicate a discarded hominin-accumulated assemblage
that was subsequently consumed by one or more carnivores
(Marean and Spencer, 1991; Marean et al., 1992). Two as-
sumptions were made concerning hominin carcass-processing
behaviors. First, it is assumed that bones were discarded in
a defleshed state, as indicated by the presence of cutmarks
on the shafts of long-bones at Pliocene (Domı́nguez-Rodrigo
et al., 2005) and Plio-Pleistocene assemblages (Bunn, 1981;
Bunn and Kroll, 1986; Domı́nguez-Rodrigo et al., 2002; Mon-
ahan, 1996; Potts, 1988; Potts and Shipman, 1981). Likewise,
it is also assumed that hominins broke limb bones to extract
marrow. This behavior is supported by fragmented shafts
and the presence of hammerstone/anvil percussion marks on
long-bones from numerous Plio-Pleistocene assemblages,
 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034
including FLK-Zinjanthropus at Olduvai Gorge, Tanzania
(Blumenschine, 1995; Blumenschine and Marean, 1993;
Bunn, 1981).
Unbroken sheep (Ovis aries) bones were purchased from
a meat distributor. Bones most commonly available included
hindlimb elements, the femur, tibia, metatarsal, and tarsals,
as well as pelves and various vertebrae. Only the long-bones
are considered in this study. Adhering flesh, fat, and cartilage
was stripped from all bones using a metal knife. Most limb
bones were broken using hammerstone and anvil percussion
(as illustrated by Binford, 1981) and the marrow was subse-
quently removed from the medullary cavities. An additional
group of limb bones were left unbroken to simulate two basic
types of sites: those with limb bones broken by hammerstone
percussion and those with unbroken limb bones. Given that
our focus is on carnivore destruction of bone, a process medi-
ated by the related properties of bone density and grease con-
tent, we do not distinguish between the two types of simulated
sites in the following analyses. The density and grease content
of cancellous limb bone portions remains constant, regardless
of how the bones are processed, and Marean and Spencer
(1991) report no significant differences between hammer-
stone-broken and unbroken assemblages.
Fig. 1. Plan of a hammerstone-broken simulated site prior to, and following, spotted hyena consumption (modified from Marean and Spencer, 1991).
2027
All simulated sites involved from one to three limb units in
addition to a vertebral section. A limb unit is defined as one
pelvis, femur, tibia, and metatarsal with adhering tarsals and
first phalanges. The vertebral sections include either a section
from the atlas to third thoracic, or the fifth and sixth lumbar
vertebrae plus the sacrum. Fig. 1 illustrates a typical simulated
site. The simulated sites were placed within enclosures at the
Berkeley Spotted Hyena Colony. As a highly effective bone-
cruncher, Crocuta provides an analogue for the more extreme
levels of carnivore modification that might occur in a hominid-
accumulated bone assemblage (Marean and Spencer, 1991).
At the time of the experiments, three groups of hyenas were
housed within three separate enclosures, which included
a compartment with a 7  10 m concrete floor connected via
a tunnel to a second 7  20 m compartment with a natural
soil base. All simulated sites were deposited in the soil-based
compartment.
The hyenas were restricted to the concrete compartment
while the sites were set up and were not fed for 24 h prior
to the experiment. All bones and bone fragments were
inscribed with a unique identification number, mapped and
photographed. One, two, or four hyenas were allowed access
to the simulated sites. Observations were made from a tower
2028 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034
adjacent to the enclosure and the experiments were terminated
when the hyenas appeared satiated, defined as no chewing or
sniffing of bones for 15 min. At this point the hyenas were
lured back into the tunnel and confined. All surviving bones
and bone fragments were re-numbered, mapped, described,
and collected for further analysis. As in previous studies
(Marean and Spencer, 1991; Marean et al., 1992), we report
the results from an analysis of 33 of these simulated site
experiments. In Table 1 we present the basic parameters for
each experimental trial.
It is important to note that the range of experimental param-
eters (i.e., 1e4 hyenas and 1e3 limb units) provides only
a partial perspective of the possible interactions between car-
nivores and discarded bone assemblages. However, the con-
trolled experimental observations do afford the opportunity
to directly observe taphonomic processes and their resultant
patterning on bone assemblages. A clear understanding of
the linkage between process and pattern provides a baseline
for extrapolating to other scenarios that may characterize the
archaeological record.
Following the experimental carnivore modification, all
bone fragments were identified according to element, side,
portion, section and percent circumference of a complete
preserved section. Bone portions include the five regions
described by Marean and Spencer (1991): proximal end, prox-
imal shaft, middle shaft, distal shaft, and distal end. These
regions were defined in a manner that recognized natural
changes in bone density along the long axis previously docu-
mented by Lyman (1984). The bone section includes the part
of the bone portion that is preserved, such as anterior/lateral.
Bone fragments that initially proved difficult to identify
were refit so that nearly all bone fragments were identified
to portion and section.
Minimum number of skeletal element (MNE) estimates
were calculated for each of the five bone portions following
the fraction summation approach described by Marean et al.
(2001), and used to determine skeletal element abundances
in several previously reported faunal assemblages (Frey and
Marean, 1999; Marean et al., 2000; Marean and Frey, 1997;
Marean and Kim, 1998). In the following analyses, we report
the relationship between differential bone portion survivorship
and bone density as it is mediated by the number of hyenas
and limb units included in each experimental trial. We use
the bone mineral density values for sheep previously calcu-
lated by Lam et al. (1998) using computed tomography
(CT). Shape-adjusted CT measurements provide the most ac-
curate bone density values (Lam et al., 2003).
Table 1
Parameters of the 33 experimental samples
1 Hyena 2 Hyenas
No. of experiments 12 13
Using 1 limb 6 7
Using 2 limbs 6 6
Using 3 limbs 0 0
Total no. of limb units 18 19
3. Experimental results
Table 2 shows the combined abundance in MNE of long-
bone portions for all elements and the levels of epiphyseal de-
letion according to the numbers of hyenas and limb units in
each experimental trial. To assess variation in bone portion
survival as a function of the number of limb units, we use
chi-square tests to compare overall portion representation for
1, 2 and 4 hyenas (Table 2). When one hyena is allowed to
modify the simulated sites, the overall bone portion survival
is equivalent regardless of the number of limb units included
in the experiment (c2 ¼ 0.268, p ¼ 0.992). In fact, as shown
in Table 2, similar non-significant results are obtained across
all of the experiments (2 hyenas: c2 ¼ 1.564, p ¼ 0.815; 4 hy-
enas: c2 ¼ 5.156, p ¼ 0.741). For 1, 2, or 4 hyenas, the addi-
tion of a second or third limb results in elevated levels of
epiphyseal and near-epiphyseal consumption to match the
input of new bones into the system. Thus, overall levels of
destruction, as measured by the relative abundance of long-
bone portions, are independent of the number of limbs. A
chi-square test also indicates that combined long-bone portion
abundances for experiments involving 1, 2, and 4 hyenas are
indistinguishable (c2 ¼ 10.06, p ¼ 0.261).
These tests indicate that the degree of epiphyseal and near-
epiphyseal deletion is independent of the number of hyenas
(i.e., from 1 to 4) and the number of limb units (i.e., from 1
to 3) included in the simulated sites. There is a slight increase
in the levels of epiphyseal deletion between 1 hyena versus 2
or 4, but this change is not significant.
MNE determinations for the three limb elements and sur-
viving portions after consumption by 1, 2, and 4 hyenas are
presented in Table 3. In Figs. 2e4 we plot the long-bone por-
tion abundance against Lam et al.’s (1998) determinations of
sheep bone density. To standardize the data, long-bone portion
abundance is reported in these figures as %MNE, scaled to the
most abundant portion. Dashed lines represent the least
squares regression with middle shaft portions included and
solid lines represent the regression without mid-shafts. We re-
port both Spearman’s and Pearson’s correlation coefficients for
the correlation between long-bone portion abundance and
bone density in Table 4. Since carnivore consumption of
bone is a density-mediated process (Brain, 1967, 1969; Marean
and Spencer, 1991; Marean et al., 1992), it comes as no sur-
prise that surviving long-bone portions correlate strongly and
significantly with bone density. However, following Faith and
Behrensmeyer (2006), we exclude the high density middle
shafts from the analysis because the they survive well regard-
less of levels of competition (Marean and Spencer, 1991) and
because bone-consuming carnivores are primarily interested
in the grease-bearing epiphyseal and near-epiphyseal portions,
especially when marrow within the medullary cavity has been
4 Hyenas previously consumed by hominins. Thus, the relative survival
8 of middle shafts is insensitive to carnivore competition; they
4 survive well whether competition is high or low. In Table 4
3 we see that long-bone portion abundance, excluding middle
1 shafts, correlates significantly with bone density only when
13
assemblages are subjected to secondary consumption by one
 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034 2029

Table 2
Combined MNE for all long-bone portions and the percentage of epiphyseal change (destroyed limb ends) for the different numbers of hyenas and limbs in each
experimental trial
1 hyena 2 hyenas 4 hyenas
1 limb 2 limbs Total 1 limb 2 limbs Total 1 limb 2 limbs 3 limbs Total
Portion Proximal end 9.4 13.25 22.65 3.7 4.3 7 2.25 5 0.9 8.15
Proximal shaft 16 27 43 12 19 30.5 9 8.5 6.5 24
Middle shaft 18.5 33.9 52.4 19.9 32.3 52.2 13.2 17 8.6 37.8
Distal shaft 14.1 24.1 37.9 7.8 17.3 25.1 5.3 11 4 18.3
Distal end 8 13 21 2 7 9 2 5 0 7
Epiphysis % change 51.67 63.54 59.58 86.43 84.31 85.87 82.29 72.22 95.00 80.58
c2 0.27a 1.56a 5.16a
Chi-square values reflect comparisons of long bone portion MNE across varying numbers of limbs in experiments with 1, 2 and 4 hyenas.
a
p > 0.50.

hyena. When two or more hyenas are involved, bone portion increasing competition counter to previous suggestions (Blu-
survivorship no longer correlates with bone density. Although menschine and Marean, 1993).
levels of bone destruction are equivalent across experimental Although the levels of long-bone destruction is comparable,
samples, the surviving limb epiphyses and near-epiphyseal given differing numbers of limbs, there is certainly a point at
shaft portions only correlate with bone density when direct which a carnivore will have eaten its fill and will no longer con-
competition for bone is absent. sume additional bone; that limit was not reached in this experi-
mental study. Nevertheless, it is reasonable to assume that
multiple carnivores are capable of consuming absolutely more
bone than a single individual. When faced with greater quanti-
4. Discussion ties of bone and/or bones of larger mammals, the destructive
impact of an individual carnivore will be much less than that
The experimental patterning of long-bone destruction pro- of a group of carnivores. Thus, an absence of competition
vides a means of interpreting the limb portions that survive implies few carnivores, lower potential for the destruction of
carnivore consumption in terms of competition intensity for elements discarded by hominins and greater preservation of
consumable bone. The experiments with captive spotted hy- patterns relating to hominin behavior. In contrast, high levels
enas indicate that when carnivores modify bone assemblages, of competition imply a much larger potential for the destruction
levels of destruction are independent (within the experimental of bones and the alteration of pre-consumption taphonomic
range) of the numbers of long-bones available for consump- signals. Under such a scenario, patterns of skeletal element rep-
tion and the degree of conspecific competition for those resentation relating to hominin behavior are likely to be severely
resources (Table 2). Levels of limb-end deletion certainly altered or destroyed.
reflect the degree of carnivore-mediated destruction of bone Concerning the relationship between carnivore competition
in a faunal assemblage, but gross levels of long-bone and the correlation between limb-end survivorship and bone
destruction (i.e., the relative number of destroyed epiphyses
and near-epiphyses) appear to be insensitive as a measure of
100
1 Hyena Ti3
Table 3
Long-bone portion MNE after consumption by 1, 2 and 4 hyenas
80 Mt3
Fe3
Element Portion MNE Ti4 Mt2
Fe2 Mt1
1 Hyena 2 Hyenas 4 Hyenas Fe4 Ti2
60
Femur Proximal end 4 0 2
Proximal shaft 14.5 9.5 8 Ti5 Mt4
Middle shaft 16.6 17.5 12.5 40 Mt5
Distal shaft 13 8.9 7.8
Distal end 3 0 0
20 Ti1 Fe1
Tibia Proximal end 4.2 1 0 Fe5
Proximal shaft 13 14.5 10
Middle shaft 20 19 13
0
Distal shaft 14.9 12.2 5.5 0.2 0.4 0.6 0.8 1 1.2 1.4
Distal end 10 6 5 Bone Density (g/cm3)
Metatarsal Proximal end 14.45 6 6.15
Fig. 2. %MNE of long-bone portion abundance vs. bone density (Lam et al.,
Proximal shaft 15.5 6.5 6
1998) following consumption by one hyena. Dashed line represents the regres-
Middle shaft 15.8 15.7 12.3
sion with mid-shafts and solid line represents the regression excluding mid-
Distal shaft 10 4 5
shafts. Fe, femur; Ti, tibia; Mt, metatarsal; 1, proximal end; 2, proximal shaft;
Distal end 8 3 2
3, middle shaft; 4, distal shaft; 5, distal end (see also Table 4).
2030 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034

100 Table 4
2 Hyenas Ti3
Correlations between long-bone portion abundance and bone density (see also
Fe3 Figs. 2e4)
80 Mt3
Ti2 Pearson Spearman

Ti4
R p Rs p
60
All portions
Fe2 1 hyena 0.811 <0.001 0.880 <0.001
Fe4
2 hyenas 0.758 0.001 0.755 0.001
40
Mt2 4 hyenas 0.753 0.001 0.727 0.002
Ti5 Mt1
Excluding middle shafts
20 Mt4 1 hyena 0.718 0.008 0.779 0.003
Mt5 2 hyenas 0.466 0.127 0.561 0.058
4 hyenas 0.459 0.134 0.506 0.093
Ti1 Fe5 Fe1
0 Significant values are in bold type.
0.2 0.4 0.6 0.8 1 1.2 1.4
Bone Density (g/cm3)

Fig. 3. %MNE of long-bone portion abundance vs. bone density (Lam et al.,
1998) following consumption by two hyenas. Dashed line represents the re-
gression with mid-shafts and solid line represents the regression excluding
mid-shafts (see also Table 4).
density, our observations provide experimental cross-valida-
tion of the naturalistic data presented previously by Faith
and Behrensmeyer (2006). A strong correlation between
epiphyseal and near-epiphyseal abundance and bone density
signals low levels of competition for food resources. The ab-
sence of such a correlation is indicative of increasing compe-
tition and potentially increased levels of destruction to other
elements in an assemblage. With the inclusion of middle
shafts, long-bone portion abundance correlates significantly
with bone density given their high density and resistance to de-
struction (Marean and Spencer, 1991; Marean et al., 1992;
Pickering et al., 2003). Marean et al. (1992) show that the low-
est density elements and portions are preferentially selected
for consumption by the spotted hyena. As a result, hyena
bone-choice correlates significantly with bone density. How-
ever, such a correlation depends on whether the hyenas have
the opportunity to choose the least dense, grease-rich elements
when feeding. Our experimental results indicate that in the
absence of competition (i.e., one hyena), this is indeed the
case and long-bone portion survivorship, excluding the middle
shafts, correlates significantly with bone density. In competi-
tive situations, however, the bone-consuming carnivores are
much less selective. With two or more hyenas present, Marean
et al. (1992) report two patterns: (1) if one hyena is dominant,
it will stand guard over the assemblage while others seek to
grab bones and run off, and (2) if the hyenas are equal in
rank, each would grab bones and run to opposite ends of the
enclosure. Based on these observations, it is apparent that un-
der direct competition, hyenas tend to go after whatever can be
readily acquired as opposed to what is least dense. When they
do not discriminate between high and low density elements,
the correlation between epiphyseal and near-epiphyseal abun-
dance and bone density becomes non-significant. Because we
have previously demonstrated that levels of bone destruction
do not vary according to the number of hyenas or limb units
included in the experimental sample, this relationship between
long-bone survivorship and bone density must be mediated by
which portions are actually selected for consumption as op-
posed to the overall levels of destruction.

5. Application
100
4 Hyenas Mt3 Ti3
Fe3 The results of these experimental studies are particularly
80 relevant to the interpretation of faunal assemblages from
Ti2
Plio-Pleistocene localities where carnivores have been impli-
Fe2
cated in altering original hominin behavioral signals. Origi-
60 Fe4
nally, the accumulations of faunal remains and stone tools in
Mt1 Mt2 East African Plio-Pleistocene sites were heralded as ‘‘living
40 Ti4
Ti5 Mt4 floors’’ (Leakey, 1971) or what Isaac (1978) termed ‘‘home
bases’’ or ‘‘central place foraging sites,’’ to which hominins
20 supposedly transported and shared a variety of foods, particu-
Fe1 Mt5 larly meat hunted from big game. Such interpretations pro-
Ti1 Fe5 vided the basis for assigning numerous modern human
0
0.2 0.4 0.6 0.8 1 1.2 1.4 behavioral characteristics to early Homo, including the emer-
Bone Density (g/cm3) gence of nuclear families and sexual division of labor (Clark,
1997; Isaac, 1978).
Fig. 4. %MNE of long-bone portion abundance vs. bone density (Lam et al.,
1998) following consumption by four hyenas. Dashed line represents the re- The traditional interpretations of the Plio-Pleistocene ar-
gression with mid-shafts and solid line represents the regression excluding chaeological record were strongly criticized by Binford
mid-shafts (see also Table 4). (1981), who questioned the degree to which hominins were
 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034 2031

responsible for the formation of the bone assemblages. Since 10

FLK-Zinj
then, archaeologists have focused on studying Plio-Pleistocene
rs = 0.506, p = 0.135
faunal remains from a taphonomic perspective in order to de- Ra1
8 r = 0.237, p = 0.510
velop better understandings of how these assemblages formed
(e.g., Binford et al., 1988; Blumenschine, 1995; Bunn, 1986;
Bunn and Kroll, 1986; Domı́nguez-Rodrigo, 1997, 1999; 6 Hu5

MAU
Isaac, 1983; Potts, 1988). Carnivores have been implicated
in the taphonomic histories of many Plio-Pleistocene assem-
4
blages (Leakey, 1971; Potts, 1988), and archaeologists con-
Mp1
tinue to argue about the contribution of hominin versus
Fe1 Mp5
carnivore agents to these bone assemblages (e.g., Blumenschine, 2 Fe5
1995; Capaldo, 1997; Domı́nguez-Rodrigo, 1999; Selvaggio, Ti1 Ra5 Ti5
1998). Hu1
0
In particular, the patterning of skeletal element representa- 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
tion and degree of carnivore involvement at the Plio-Pleisto- Bone Density (g/cm3)
cene sites of FLKN levels 1-2 and FLK-Zinjanthropus in
Fig. 5. Long-bone end MAU vs. bone density of wildebeest (Lam et al., 1999)
Olduvai Gorge, Tanzania, have been the source of continuous for size 3 bovids at FLK-Zinjanthropus (Potts, 1988). Mp, metapodial.
investigation and debate (Binford, 1981, 1986, 1988;
Blumenschine, 1995; Bunn, 1986; Bunn and Kroll, 1986,
1987, 1988; Capaldo, 1997; Domı́nguez-Rodrigo and Barba, experimental results under competitive scenarios (rs ¼ 0.506,
2006; Faith and Behrensmeyer, 2006; Marean et al., 1992; p ¼ 0.135; r ¼ 0.237, p ¼ 0.510). In the following discussion,
Potts, 1987, 1988; Selvaggio, 1994). Although it is not we consider the assemblages recovered from FLKN levels 1e2.
possible to identify the species of carnivore responsible for As with FLK-Zinjanthropus, the assemblage from FLKN
modifying these assemblages, damage patterns (e.g., destruc- levels 1e2 is dominated by appendicular elements. This has
tion of long-bone epiphyses (Blumenschine and Marean, been interpreted as indicating hominin-transportation of meaty
1993; Bunn, 1986; Domı́nguez-Rodrigo and Barba, 2006)) in- limb elements (Bunn, 1986). However, Marean et al. (1992)
dicate specialized bone-crunching carnivores, such as hyenas. provide a cautionary note, demonstrating that this pattern of
Fossil hyenas recovered from Beds I and II at Olduvai Gorge skeletal element representation could result from carnivore de-
include C. ultra and Hyaena sp. (Werdelin and Lewis, 2005; struction of axial elements rather than hominin limb transport.
Margaret E. Lewis, personal communication, 12/22/2006). Based on our experimental findings, we can re-examine the
We assume that these carnivores would have had preferences levels of carnivore competition for the hominin-discarded as-
for skeletal elements and long-bone portions similar to those semblages at FLKN levels 1e2. In Fig. 6, we plot limb-end
of modern C. crocuta, since these preferences are related to abundances provided by Bunn (1986) for size categories 3
the nutritional value (i.e., grease content) and density of and 4 mammals, the majority of which are bovids, against
bone. The assemblages from FLK-Zinjanthropus and FLKN the bone density values for wildebeest reported by Lam
levels 1e2 are characterized by an abundance of limbs and et al. (1999). A very strong and positive correlation is apparent
very low frequencies of axial elements. The relative rarity of (rs ¼ 0.720, p ¼ 0.008; r ¼ 0.675, p ¼ 0.016), suggesting low
axial remains at these sites may reflect hominin transport of
appendicular elements, carnivore destruction of axial ele-
ments, or a combination of both (Marean et al., 1992). Faith 50
FLKN Levels 1-2
and Behrensmeyer (2006) suggest that a poor correlation be-
rs = 0.720, p = 0.008
tween epiphyseal abundance and bone density at FLK-Zinjan- 40 Ti5
r = 0.675, p = 0.016
thropus indicates intense carnivore competition and high
levels of carnivore modification. Thus, our ability to make in- Hu5
ferences about hominin behavior based on skeletal element 30 Ra1 Mt1
MNE

abundances at FLK-Zinjanthropus is limited. The experimen- Mt5 Mc1

Mc5
tal results from our study support this interpretation of the 20
FLK-Zinjanthropus archaeofauna. Here, we plot limb-end Ra5
abundance of size 3 bovids provided by Potts (1988) against Ti1
Lam et al.’s (1999) wildebeest (Connochaetes taurinus) bone 10
Fe5 Fe1
density values (Fig. 5). Bone density values for metacarpals Hu1

and metatarsals are averaged as Potts reports these elements

0
as metapodials. Limb-end frequency is reported in minimal 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
animal units (MAU) after Binford (1984) to standardize the Bone Density (g/cm3)
abundance of metapodials relative to the other elements. The Fig. 6. Long-bone end MNE vs. bone density of wildebeest (Lam et al., 1999)
strength of the correlation between limb-end abundance and for size 3 and 4 mammals at FLKN levels 1e2 (Bunn, 1986). Ra, radius; Mc,
bone density is not significant, which is consistent with the metacarpal.
2032 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034
levels of carnivore competition. Bunn’s report on the FLKN
levels 1e2 assemblage provides highly accurate MNE deter-
minations because he included limb shafts in the counts (see
our Table 5). This allows us to calculate the total level of
epiphyseal destruction by carnivores and other agents at
28.8%, a value much lower than the experimental values for
1e4 captive hyenas.
Because the processes of hominin-discard and secondary
carnivore consumption contributing to the formation of the
FLKN levels 1e2 archaeofauna took place over an unknown
length of time, our measurements reflect a time-averaged re-
gime of carnivore competition that, overall, appears to be quite
low. Had axial remains initially been deposited at this site, it is
highly unlikely that such a low number of carnivores, perhaps
solitary individuals acting sequentially on the assemblage,
would have destroyed large quantities of axial elements from
size 3e4 mammals. In their study of carnivore destruction
of bone on the Amboseli landscape, Faith and Behrensmeyer
(2006) show that only under extremely high carnivore densi-
ties and elevated levels of competition are the axial remains
of size 2 and 3 ungulates thoroughly destroyed. A low estimate
of limb-end deletion in tandem with the strong correlation be-
tween limb-end abundance and bone density suggests that the
FLKN levels 1e2 assemblage was subjected to minimal levels
of carnivore attrition and that the faunal remains recovered
from the assemblage likely retain a strong signal of hominin
behavioral patterns relative to taphonomic overprinting caused
by carnivore activity.
We conclude that the skeletal element representation at
FLKN levels 1e2 is consistent with a pattern of hominin discard
of meaty appendicular elements rather than carnivore destruc-
tion of axial elements with consequent preferential survival of
appendicular elements. These findings contrast with the results
of an equivalent analysis of FLK-Zinjanthropus, which appears
to have been modified by carnivores under high levels of com-
petition (Faith and Behrensmeyer, 2006). It is possible that the
Table 5
Long-bone abundances of size 3/4 mammals at FLKN levels 1e2 as deter-
mined by Bunn (1986)
Element
Humerus Proximal
Distal
Shafts included
Radius Proximal
Distal
Shafts included
Metacarpal Proximal
Distal
Shafts included
Femur Proximal
Distal
Shafts included
Tibia Proximal
Distal
Shafts included
Metatarsal Proximal
Distal
Shafts included
taphonomic history of FLK-Zinjanthropus may be atypical
relative to other contemporaneous sites. FLK-Zinjanthropus
has been the focus of numerous analyses, resulting in a wide
range of interpretations for the implications of hominin behav-
ioral patterns (Binford, 1981, 1986, 1988; Blumenschine,
1995; Bunn, 1986; Bunn and Kroll, 1986, 1988; Capaldo,
1997; Domı́nguez-Rodrigo and Barba, 2006; Faith and Behren-
smeyer, 2006; Potts, 1988; Selvaggio, 1994, 1998). The diver-
gent taphonomic histories of FLK-Zinjanthropus and FLKN
levels 1e2 stress the need for caution when making inferences
concerning the evolutionary history of hominin foraging strate-
gies based on the study of a single, unusually rich put possibly
atypical archaeofaunal assemblage.
6. Conclusions
Inferring hominin behavior from the patterns evident in the
fossil record requires an understanding of the taphonomic pro-
cesses that have mediated skeletal element representation. It
has long been recognized that a skeletal element’s physical
characteristics, most notably bone density, play a central role
in the differential survivorship of bones and portions of bones
(Brain, 1967, 1969). This is particularly relevant to carnivore-
mediated attrition, which has played a significant role in
patterning the faunal remains from numerous assemblages of
paleoanthropological interest. A well-developed understanding
of how carnivore-mediated taphonomic signals relate to varia-
tion in ecology, competition for nutrient resources and carni-
vore behavioral patterns can significantly enhance our ability
to infer hominin behavior and general paleoecological condi-
tions from archaeofaunal and paleontological assemblages al-
tered by carnivores (Blumenschine and Marean, 1993; Gifford,
1981).
In this study we have re-analyzed previous results from
controlled experimental observations of spotted hyenas in or-
der to examine the relationships between carnivore competi-
tion, bone destruction, and bone density. The overall levels
of carnivore-mediated long-bone destruction in simulated
hominid-discarded assemblages is unaffected by the number
of hyenas (1, 2 or 4) permitted to consume the bone, as well
MNE as the quantity of bone discarded at each site. Thus an evalu-
ation of overall levels of epiphyseal deletion at a faunal assem-
7
33 blage does not provide insight into the number of carnivores
33 involved in modifying the bone, at least within the experimen-
30 tal range. The experimental results also indicate that surviving
17 long-bone portions correlate strongly with bone density, pri-
30
marily because high density middle shafts survive carnivore
25
23 consumption regardless of levels of competition. The consis-
25 tently high survival of mid-shaft portions act as leverage
9 points, generating highly significant positive correlations in
9 an otherwise uncorrelated scatter of points. It follows that in
33
evaluating levels of conspecific competition for bone, middle
13
41 shafts should be excluded from the statistical analysis because
41 of their high survival potential. In support of previous observa-
30 tions based on the modern landscape bone assemblage of
25 Amboseli Park, Kenya (Faith and Behrensmeyer, 2006) the
30
experimental results indicate that the correlation between
 J.T. Faith et al. / Journal of Archaeological Science 34 (2007) 2025e2034
long-bone epiphyseal and near-epiphyseal abundance and
bone density does provide a measure of overall levels of
carnivore competition. Strong correlations signal low intensity
competition because bone-consuming carnivores can preferen-
tially select low density elements for consumption. This signal
also implies a lower potential for overall destruction of other
elements (e.g., axial elements) because few carnivores are
competing for the skeletal remains. Such a pattern was only
evident in experimental trials involving a single hyena. A
non-significant correlation, in contrast, signals elevated levels
of direct competition for bone. This provides evidence for the
involvement of multiple carnivores in modifying an assem-
blage. In such cases, behavioral signals from previous hominin
involvement with skeletal element representation are likely to
be overprinted or destroyed.
Acknowledgements
We are grateful to Donald Grayson, Margaret Lewis and the
anonymous reviewers for helpful comments on previous ver-
sions of this manuscript. JTF thanks the National Science
Foundation for supporting this research under a Graduate Re-
search Fellowship. CWM thanks the helpful staff at the Berke-
ley Spotted Hyena Colony, Berkeley, California. Laurence
Frank was particularly helpful during the research with the hy-
enas, and Stephen Glickman consented to the experiments.
The captive colony was supported by the National Institute
of Health grant No. 5R01 MH 39917 to Stephen Glickman
during the time of these studies. CWM also thanks Hope
Williams for assisting with the figure production, and the
ISSR staff for technical support.
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