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Journal of Human Evolution xx (2007) 1e7

News and Views

Carnivore tooth-marks, microbial bioerosion, and the invalidation of

Domı́nguez-Rodrigo and Barba’s (2006) test of Oldowan hominin
scavenging behavior
Robert J. Blumenschine*, Kari A. Prassack, C. David Kreger, Michael C. Pante
Center for Human Evolutionary Studies, Department of Anthropology, Rutgers University, 131 George St., New Brunswick, NJ 08901-1414, USA.
Received 27 September 2006; accepted 8 January 2007

Keywords: Taphonomy; Olduvai Gorge

Introduction and in some cases large quantities of flesh on unattended natural

The title of a recent paper by Domı́nguez-Rodrigo and
Barba (2006; ‘‘New estimates of tooth mark and percussion
mark frequencies at the FLK Zinj site: the carnivore-homi-
nid-carnivore hypothesis falsified’’) trumpets the partial reso-
lution of a long-standing debate on the role of hunting and
scavenging in Oldowan hominin adaptation. It proclaims that
larger mammal carcasses represented by fossil bones from
the Zinjanthropus level at FLK, Olduvai Gorge, were not con-
sumed first by flesh-eating carnivores followed by scavenging
hominins, and then bone-crunching carnivores. In short, it
claims that Oldowan hominins did not scavenge carcasses con-
sumed partially by other carnivores.
In the abstract, Domı́nguez-Rodrigo and Barba (2006) ac-
knowledge the title’s overstatement by claiming they have fal-
sified the passive scavenging hypothesis, one variation of the
‘‘carnivore-hominin-carnivore’’ sequence of carcass access.
Other hypothesized examples of secondary access to carcasses
by hominins d with or without tertiary access by carnivores d
range in food yield and danger from ‘‘marginal scavenging’’
of negligible, abandoned remains (Binford, 1981), to ‘‘power
scavenging’’ (Bunn, 2001) or confrontational scavenging,
where hominins usurp largely complete carcasses from preda-
tors. Passive scavenging affords hominins intermediate food
yields with minimal predation risk, providing marrow, the brain,
* Corresponding author.
E-mail address: rjb@rci.rutgers.edu (R.J. Blumenschine).
deaths or on abandoned felid kills (e.g., Blumenschine, 1986a,
1987; Cavallo and Blumenschine, 1989). It has withstood tests,
including those based on Blumenschine’s estimates of the fre-
quency of tooth-marked and percussion-marked long bones of
larger mammals at FLK Zinjanthropus (Blumenschine, 1995;
Capaldo, 1997; Selvaggio, 1998). Domı́nguez-Rodrigo and
Barba (2006) provide revised estimates of tooth-mark frequencies
at FLK Zinjanthropus that suggest to them that Oldowan homi-
nins had ‘‘early access to fleshed carcasses,’’ with hunting, in their
view, being more likely than confrontational scavenging. Here,
we demonstrate that the revised estimate is unsupported and its
implication unwarranted.
Carnivore tooth-marking and hominin paleoecology
Blumenschine (1988) developed a methodology for using
carnivore tooth-marking to assess the mode of carcass acquisi-
tion by hominins. This methodology has been expanded on by
others (Blumenschine and Marean, 1993; Selvaggio, 1994,
1998; Capaldo, 1995, 1997; Lupo, 1995; Marean and Kim,
1998; Marean et al., 2000), and is used by Domı́nguez-
Rodrigo and Barba (2006). The methodology uses controlled
observations of stone-tool butchery and carnivore feeding in
different sequences to interpret frequencies of tooth-marked,
percussion-marked, and cut-marked fossil bones.
Domı́nguez-Rodrigo and Barba’s core claim is that Blu-
menschine (1995) overestimated the number of long bone
midshaft specimens that bear carnivore tooth marks in the

0047-2484/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jhevol.2007.01.011

Please cite this article in press as: Robert J. Blumenschine et al., Carnivore tooth-marks, microbial bioerosion, and the invalidation of Domı́nguez-Rodrigo and
Barba’s (2006) test of Oldowan hominin scavenging behavior, J Hum Evol (2007), doi:10.1016/j.jhevol.2007.01.011
 ARTICLE IN PRESS
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2 R.J. Blumenschine et al. / Journal of Human Evolution xx (2007) 1e7
FLK Zinjanthropus assemblage due to his alleged confusion of
these with what Domı́nguez-Rodrigo and Barba (2006) label
as ‘‘biochemical marks.’’ They claim to have recognized and
broken a new equifinality between traces on bone produced
by microorganisms that degrade bone of dead animals, and
tooth marks produced by vertebrate carnivore feeding. Domı́-
nguez-Rodrigo and Barba (2006) produce a revised estimate of
tooth-marked midshaft frequencies in the FLK Zinjanthropus
assemblage (11%) that is more than five times lower than Blu-
menschine’s (58%).
Their new estimate of the frequency of tooth-marked mid-
shafts lies within the 95% confidence intervals (3.5% e
17.5%) of the mean assemblage value produced when modern
spotted hyenas have access only to grease in long bones that
had been butchered and hammerstone-broken for marrow (the
‘‘hominin-carnivore’’ model; Blumenschine, 1988, 1995;
Capaldo, 1997, 1998; Marean and Kim, 1998; Marean et al.,
2000). This contrasts with the implications of Blumenschine’s
original estimate (58%), which lies well above the upper
confidence interval of the ‘‘hominin-carnivore’’ model. Blumen-
schine attributed the high frequency of tooth-marked midshafts
at FLK Zinjanthropus to felid defleshing of whole bones later
scavenged passively by hominins. Selvaggio’s (1994, 1998)
neotaphonomic work demonstrates such high rates of midshaft
tooth-marking from carnivore defleshing in her ‘‘carnivore-
hominin’’ (47%) and ‘‘carnivore-hominin-carnivore’’ (54%)
sequences of carcass access. Since epiphyseal deletion by carni-
vores occurred at FLK Zinjanthropus (e.g., Blumenschine and
Marean, 1993), the latter value is most appropriately compared
to d and is less than four percentage points lower than d the
proportion of tooth-marked midshaft specimens at FLK
Zinjanthropus obtained by Blumenschine.
Readers familiar with this result would recognize the error
in Domı́nguez-Rodrigo and Barba’s claim that the frequencies
of midshaft tooth-marking obtained by Blumenschine for FLK
Zinjanthropus ‘‘stood out as an oddity never before replicated
experimentally’’ (Domı́nguez-Rodrigo and Barba, 2006:171),
and, again, that ‘‘the frequencies..... fell outside the ranges
of variation for all experimental assemblages’’ (Domı́nguez-
Rodrigo and Barba, 2006:180). Their mistake arises from an
oft-repeated (e.g., Domı́nguez-Rodrigo, 2002:30; Domı́-
nguez-Rodrigo and Pickering, 2003:279) confusion of tooth-
mark frequencies obtained in Selvaggio’s ‘‘carnivore-first’’
neotaphonomic models with Blumenschine’s (1988) ‘‘carni-
vore-only’’ model, in which carnivores alone are responsible
for extracting all edible tissues from long bones. For example,
in referring to their low (9.7%) estimated frequency of tooth-
marked midshafts on ‘‘large-sized carcasses’’ from FLK Zin-
janthropus, Domı́nguez-Rodrigo and Barba (2006:179) state
that this value ‘‘is far from the expected percentage (86.5%)
for a carnivore-first scenario required by the carnivore-homi-
nid-carnivore hypothesis’’ (emphasis added). The 86.5% value
they cite was obtained by Blumenschine for wildebeest-sized
animals in the carnivore-only model, which, by definition, ex-
cludes hominins and does not serve as a neotaphonomic ex-
pectation for rates of tooth-marking associated with hominin
scavenging.
Please cite this article in press as: Robert J. Blumenschine et al., Carnivore tooth-marks, microbial bioerosion, and the invalidation of Domı́nguez-Rodrigo and
Barba’s (2006) test of Oldowan hominin scavenging behavior, J Hum Evol (2007), doi:10.1016/j.jhevol.2007.01.011
Carnivore tooth-marking and microbial bioerosion
The term ‘‘biochemical mark’’ introduced by Domı́nguez-
Rodrigo and Barba (2006) is known as ‘‘bioerosion’’ in the pa-
leontological, archaeological, and forensic literatures. Coined
four decades ago by Neumann (1966), bioerosion is the chem-
ical or mechanical modification of hard substances by micro-
organisms, marine invertebrates, insects, and vertebrates (e.g.,
Neumann, 1966; Davis, 1997). The term is used predomi-
nantly to describe modifications made by microbes and marine
invertebrates. The first description of microbial bioerosion on
vertebrate tissues is attributed to Wedl (1864).
Most of the modern literature on bioerosion of vertebrate
bone describes microscopic modification of its internal histo-
logical structure by soil and endogenous gut microorganisms
(fungi, algae, bacteria, amoebic protozoa), some or all of
which bioerode bone while demineralizing it to consume col-
lagen (e.g., Child et al., 1993; Hedges et al., 1995; Turner-
Walker and Syversen, 2002). Fewer studies describe microbial
bioerosion on bone surfaces, and only Domı́nguez-Rodrigo
and Barba (2006) claim that fungal bioerosion is similar to
carnivore tooth-marking.
Archaeological and paleontological studies of microbial bi-
oerosion include descriptions of bioeroded fossils and subfos-
sils, and experimental work involving the inoculation of bone
by known microorganisms and controlled exposure of bone to
various soil and aqueous environments (e.g., Marchiafava
et al., 1974; Hackett, 1981; Ascenzi and Silvestrini, 1984; Pie-
penbrink, 1986; Child et al., 1993; Child, 1995; Dye et al.,
1995; Hedges et al., 1995; Bell et al., 1996; Nicholson,
1996, 1998; Davis, 1997; Jackes et al., 2001; Trueman and
Martill, 2002; Jans et al., 2004). These studies show that mi-
crobial bioerosion can develop within several days of death
(Davis, 1997), and can occur on a majority of bones in exper-
imental and fossil assemblages (e.g., Trueman and Martill,
2002).
Development of microbial bioerosion is influenced by a va-
riety of environmental factors (e.g., Bell et al., 1996; Nichol-
son, 1996, 1998; Davis, 1997), mode of death and scavenger
activity (Bell et al., 1996; Dent et al., 2004), and characteris-
tics of bone. Microbes invade cancellous bone more readily
than compact bone (Sharmin et al., 2003), and they bioerode
porous, epiphyseal margins more extensively than compact
bone (Nicholson, 1998). The vascularized surface of mamma-
lian cortical bone is colonized by microbes more readily than
the denser cortical bone of birds (Nicholson, 1996).
Most of this large literature on microbial bioerosion is ac-
cessible and widely cross-referenced. Little of it is cited by
Domı́nguez-Rodrigo and Barba (2006), such that their work
on the subject is largely uninformed by a long history of exper-
imental and paleontological studies.
Domı́nguez-Rodrigo and Barba claim tooth mark mimicry
by microbial bioerosion on the basis of ‘‘preliminary’’ results
of a study involving 20 equid and bovid bones that had been
placed in a box and inspected occasionally for about two
years. The authors claim repeatedly that this provides an ‘‘ex-
perimentally-derived referential framework’’ for identifying
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R.J. Blumenschine et al. / Journal of Human Evolution xx (2007) 1e7
marks produced by fungi, but they provide virtually no details
on experimental protocol. Aside from learning that the bones
had been defleshed and their marrow removed, the reader is
not informed about the source and initial surface condition
of the bones, the conditions to which the box of bones was ex-
posed, and the manner in which staining and bone erosion
were linked securely to fungi as opposed to the many other or-
ganisms known to bioerode bone. Also lacking are control sets
of bone, such as those on which fungal attack was inhibited or
attack by other organisms was promoted.
Many studies of bioerosion report bone staining of various
colors by microorganisms (e.g., Piepenbrink, 1986; Nicholson,
1996, 1998). Domı́nguez-Rodrigo and Barba (2006:176) note
that ‘‘a few’’ of the stained areas on the ‘‘experimental’’ bones
showed ‘‘micro-flaking’’ or ‘‘exfoliation,’’ ‘‘creating pits or
scores that could be mistaken for a tooth mark in the absence
of a distinct outline coloring.’’ Domı́nguez-Rodrigo and Bar-
ba’s (2006) attribution of staining on some FLK Zinjanthropus
fossils to microorganisms seems reasonable. Blumenschine
noted staining associated with marks during his analysis of
the FLK Zinjanthropus assemblage, attributing it to root-
etching (e.g., Binford, 1981). Some marks attributed by
Blumenschine to carnivore teeth may have also been stained.
However, because some fungi may colonize bone at sites mod-
ified previously by other agents (e.g., Nicholson, 1998), the
crushed interior surfaces of carnivore tooth marks may be
especially susceptible to microbial attack. We have initiated
experiments to test this proposition.
No prior researcher has described patterns of microbial bi-
oerosion that could be interpreted as scores or pits of a size
and configuration similar to those produced by carnivore teeth.
Most microbial bioerosion is referred to as ‘‘tunnels’’ [more
technically ‘‘microscopical focal destruction’’ (Hackett,
1981)]. Four of the five recognized types of microscopic tun-
nels refer to bioerosion within bone (Davis, 1997). The fifth
type, labeled Hackett tunnels by Davis (1997), develops on
bone surfaces and may be more appropriately described as
channels. Microscopic pitting is also reported, although less
commonly. Individual surface channels and pits have widths
typically around 10 microns, with examples up to 100 microns
(Figs. 1a, c; Davis, 1997). Nicholson (1996) reports lengths of
surface bioerosion channels between 0.5 and 2.0 mm, and Da-
vis (1997) illustrates three specimens with surface channels no
longer than about 1 mm. Tooth pits and scores of large mam-
malian carnivores and crocodiles are typically much larger, by
up to one order of magnitude (e.g., Domı́nguez-Rodrigo and
Piqueras, 2003; Njau and Blumenschine, 2006). Even smaller
carnivores such as domestic cats, skunks, and genets produce
pits with diameters  1 mm and scores with similar widths
(Prassack, 2006). Unfortunately, Domı́nguez-Rodrigo and
Barba (2006) provide scale for only one of 21 photographs
of alleged bioerosion from their modern and fossil samples;
the scaled specimen they depict has a score with a maximum
width 1 mm, typical of tooth marks and outside the pub-
lished range of known bioerosion channels.
Domı́nguez-Rodrigo and Barba’s (2006) inattention to
scale leads to their misuse of the following quote from
Please cite this article in press as: Robert J. Blumenschine et al., Carnivore tooth-marks, microbial bioerosion, and the invalidation of Domı́nguez-Rodrigo and
Barba’s (2006) test of Oldowan hominin scavenging behavior, J Hum Evol (2007), doi:10.1016/j.jhevol.2007.01.011
3
Sharmin et al. (2003:19): ‘‘The fungal isolates exhausted the
architecture of compact cortical bones by creating irregular
cratered surfaces.’’ Domı́nguez-Rodrigo and Barba (2006:
175) embellish this quote by adding that the irregular surfaces
were cratered ‘‘and grooved.’’ While ‘‘cratered and grooved
surfaces’’ might evoke those pitted and scored by carnivore
teeth, Sharmin et al. (2003) are referring to internal micro-
scopic damage that was detected on 4e6 micron thin-sections,
illustrated with photographs at 150 magnifications.
The anatomical patterning and morphology of microbial bi-
oerosion on bone surfaces are also not evocative of carnivore
tooth-marking. Surface clusters of bioerosion channels and
pits are often extensive (Fig. 1c), unlike isolated tooth pits
or scores (Figs. 1d, e). Bioerosion channels are described as
forming a ‘‘meandering pattern,’’ a ‘‘branched meandering
pattern’’ (Davis, 1997), or a ‘‘net-like (so-called root) pattern’’
(Tappen, 1994:669), and producing a ‘‘rippled or scalloped’’
texture (Nicholson, 1996) that does not mimic tooth-marked
surfaces. Further, the internal morphology of microbial marks
is smooth or composed of a cluster of micro-pits that is dis-
tinctive from the crushing that characterizes carnivore tooth
marks (compare Figs. 1a, c to Figs. 1d, e).
The criteria Domı́nguez-Rodrigo and Barba (2006) use to
distinguish surface marks produced by microorganisms and
carnivore teeth in the absence of staining on the FLK Zinjan-
thropus bones are described qualitatively only. As a result, the
reader can evaluate neither the statistical strength of the
distinctions nor replicate the results. Further, they provide no
information on the source of tooth-marked or ‘‘biochemically-
marked’’ bones used to establish these criteria. Are the distinc-
tions based on the ‘‘few’’ ‘‘biochemically-marked’’ bones they
have produced ‘‘experimentally,’’ and are these compared to
a sample of modern bones tooth-marked by carnivores under
controlled conditions? Or, given that these differences are illus-
trated only by examples of alleged ‘‘biochemical marks’’ on fos-
sil bones from FLK Zinjanthropus, are they using the presence
and absence of staining to differentiate the two kinds of marks in
the assemblage? The authors warn that their ‘‘results must be
treated with caution [because] experimentally-derived models
for biochemical damage remain incomplete’’ (Domı́nguez-
Rodrigo and Barba, 2006: 175). We agree. Nonetheless, they
conclude that ‘‘the carnivore-hominin-carnivore hypothesis,
the epitome of the passive scavenging models, has been falsified
by the present study.’’ (Domı́nguez-Rodrigo and Barba, 2006:
189). This claim is at best premature and most likely wrong.
Domı́nguez-Rodrigo and Barba (2006) may have underesti-
mated the frequency of tooth-marking at FLK Zinjanthropus
because many tooth-marked specimens may have been bio-
eroded microbially as well. This possibility is enhanced by re-
ports that bioerosion develops more readily on bone that lacks
flesh or fat (e.g., Nicholson, 1996, 1998; Davis, 1997), a cir-
cumstance that characterized the long bone fragments in the
assemblage. Domı́nguez-Rodrigo and Barba’s (2006) Table 1
reports that of the 255 specimens in a shared sample that Blu-
menschine originally identified as being tooth-marked, they
identified 61 as tooth-marked and 190 as biochemically
marked, leaving four undiagnosed specimens. Apparently,
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Fig. 1. Examples of known and inferred microbial and insect bioerosion, mammalian carnivore tooth-marking, and hammerstone percussion marking on the sur-
faces of modern (a, d) and fossil bones from Bed I and Lower Bed II, Olduvai Gorge (b, c, e, f). (a) Microbial bioerosion in the form of a ca. 2 mm long oblong
depression developed on a chicken ulna that had been submerged in a sealed container filled with fresh water from a Typha marsh for 11 months. The bioerosion,
stained black interiorly, had been covered by a black film thought to be algae and had a patch of white mold (under which no bioerosion was detected) immediately
adjacent to it. The oblong depression is formed by a densely-packed cluster of pits each < 100 mm in diameter, forming a dimpled interior surface unlike the
crushed surfaces of carnivore tooth marks. Several of these pits are also found to the left of the oblong depression. (b) Channeling on the midshaft of a me-
dium-sized bovid tibia. The density and multi-directionality of the channels, and the length, width, and smooth internal morphology of individual channels are
similar to damage attributed to termites by Fejfar and Kaiser (2005, Fig. 8). (c) Branched-meandering white staining on the metapodial midshaft of a small bovid
likely produced by microorganisms. Widespread bioerosion in the form of pits < 100 mm in diameter are found within the staining, coalescing into discontinuous
channeling in some places. On other examples, such pitting and channeling occur outside of stained areas. Linear striations, most likely caused by trampling,
extend diagonally across the stained area. (d) Modern carnivore tooth pit with emanating score on a medium-sized mammal long bone midshaft fragment, showing
the diagnostic crushed interior surface lacking in all examples of microbial bioerosion we have seen. (e) Two inferred carnivore tooth pits on a fossil cranial frag-
ment. One pit has an emanating score. Both pits and the score show the crushing seen in the modern example in (d). Smaller, less conspicuous carnivore tooth pits
and scores are diagnosed using the same criterion (Blumenschine et al., 1996). (f) Inferred and very conspicuous percussion marking on a medium-sized bovid
metapodial midshaft. The interior surfaces of individual marks are not crushed, but have parallel sets of microstriations that distinguish it from carnivore tooth-
marking and non-vertebrate bioerosion. Length of scale bars ¼ 2 mm.

Please cite this article in press as: Robert J. Blumenschine et al., Carnivore tooth-marks, microbial bioerosion, and the invalidation of Domı́nguez-Rodrigo and
Barba’s (2006) test of Oldowan hominin scavenging behavior, J Hum Evol (2007), doi:10.1016/j.jhevol.2007.01.011
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R.J. Blumenschine et al. / Journal of Human Evolution xx (2007) 1e7
they did not identify tooth-marking if a specimen bore alleged
‘‘biochemical marks,’’ a standard that will always underesti-
mate tooth-mark frequencies.
Domı́nguez-Rodrigo and Barba’s (2006) underestimation of
tooth-mark frequencies is also suspected on the basis of their
substantially lower estimate of percussion mark frequencies
compared to Blumenschine’s (1995). They attribute this dis-
crepancy to the erroneous supposition that Blumenschine
counted percussion notches as percussion marks. Yet, Blumen-
schine and Selvaggio (1988, 1991) introduced the distinction
between the fracture patterns and surface marks produced by
hammerstone percussion, and Blumenschine (1995, his
Tables 3a and 4) reported their occurrences on the FLK Zinjan-
thropus assemblage separately. Instead, Domı́nguez-Rodrigo
and Barba (2006) have clearly missed a number of percussion
marks, which, unlike the prominent example in Fig. 1f, can be
very inconspicuous. Our contention is supported by Capaldo’s
virtually identical estimate of percussion mark frequencies on
long bone midshafts to that obtained by Blumenschine in his
independent analysis of FLK Zinjanthropus (Blumenschine
et al., in press). Given that many bones bearing carnivore feed-
ing traces are tooth-marked inconspicuously (e.g., Blumen-
schine et al., 1996), Domı́nguez-Rodrigo and Barba’s (2006)
failure to detect many percussion marks on FLK Zinjanthropus
bones likely applies to tooth marks as well.
Domı́nguez-Rodrigo and Barba (2006:179) misuse a result
of Cushing’s (2002; Table 6.1) analysis of bones recovered by
Blumenschine and Masao’s ongoing excavations at Olduvai by
claiming that ‘‘biochemical marks are present on bones from
almost half of the occurrences sampled in the Gorge.’’ How-
ever, Cushing never uses the term ‘‘biochemical marks,’’ refer-
ring instead to ‘‘chemical pitting’’ she attributes to carnivore
digestion and/or soil chemistry. Further, the proportion of
bones she identified as chemically pitted is only 1.4%.
Carnivore tooth-marking and passive scavenging
Domı́nguez-Rodrigo and Barba (2006) assume implicitly
that passive scavenging will always be associated with fre-
quencies of tooth-marked long bone midshaft fragments that
are significantly higher than those associated with confronta-
tional scavenging or hunting by hominins. This assumption
is consistent with Selvaggio’s ‘‘carnivore-first’’ results. How-
ever, it is contradicted by reports that purport to show that fe-
lids inflict few tooth marks on long bone midshafts during
defleshing, and that hammerstone-broken bones defleshed pre-
viously by felids yield a frequency of tooth-marked midshaft
fragments that lies within the range expected for ‘‘hominin-
first’’ models. Oddly, Domı́nguez-Rodrigo (1999; Domı́-
nguez-Rodrigo et al., in press, based on a paper delivered
orally in 2004) authored these observations and interpreta-
tions, but these works are neither cited nor discussed by Dom-
ı́nguez-Rodrigo and Barba (2006). The reader is therefore left
with a dilemma: dismiss either Domı́nguez-Rodrigo’s observa-
tions and interpretations of tooth-marking by modern felids, or
Domı́nguez-Rodrigo and Barba’s (2006) conclusion that the
Please cite this article in press as: Robert J. Blumenschine et al., Carnivore tooth-marks, microbial bioerosion, and the invalidation of Domı́nguez-Rodrigo and
Barba’s (2006) test of Oldowan hominin scavenging behavior, J Hum Evol (2007), doi:10.1016/j.jhevol.2007.01.011
5
‘‘carnivore-hominid-carnivore hypothesis’’ for Oldowan hom-
inins has been falsified.
Domı́nguez-Rodrigo and Barba (2006) make another un-
warranted implicit assumption. They assume that passive scav-
enging will always be associated with hominin access to
largely defleshed bones remaining with only flesh scraps.
Yet, an established body of literature hypothesizes a wide
range of passive scavenging opportunities for hominins that
provide large quantities of flesh, including complete carcasses
(e.g., Blumenschine, 1986a; Haynes, 1988; Cavallo and Blu-
menschine, 1989; Marean, 1989; Capaldo and Peters, 1995;
Werdelin and Lewis, 2005). For example, hominins scaveng-
ing natural deaths passively would access fully-fleshed long
bones lacking midshaft tooth-marking, mimicking ‘‘hominin-
first’’ scenarios. Passive scavenging of carcasses defleshed
only partially by carnivores would yield tooth marks on only
those bones fed on relatively early in the carcass consumption
sequence (Blumenschine, 1986b). The result would be lower
overall rates of tooth-marking than those associated with pas-
sive scavenging of fully defleshed kills, an expectation sup-
ported by Pante’s (2006) observations of a carcass partially
defleshed by a leopard. In this study, only the hind limbs
had been defleshed, such that after hammerstone breakage of
all scavenged limb bones, the overall incidence of tooth-
marking on midshaft fragments was only 20%. These findings
underscore the need to study tooth-marking associated with
a fuller range of modern scavenging opportunities. They also
reinforce the erroneous ‘‘falsification’’ of the ‘‘carnivore-
hominid-carnivore’’ hypothesis by Domı́nguez-Rodrigo and
Barba (2006).
Conclusions
We have demonstrated that Domı́nguez-Rodrigo and Bar-
ba’s (2006) falsification of the ‘‘carnivore-hominid-carnivore’’
hypothesis is an exaggerated claim based on pseudo-experi-
mental procedures, limited reading of relevant literatures,
and unreplicable observations of modern and fossil bones.
Their case for microbes bioeroding bone in ways that mimic
carnivore tooth-marking is unsubstantiated, as are the criteria
they use for breaking this alleged equifinality. As such, their
revised estimate of the frequency of tooth-marked bones at
FLK Zinjanthropus is invalid. Given the wide variety of pas-
sive scavenging opportunities Oldowan hominins likely en-
countered, their blanket rejection of this type in favor of
confrontational scavenging and especially hunting is a general-
ization unsupported by what we know or can expect about
feeding traces associated with passive scavenging. In sum,
they have failed to falsify passive scavenging from felid kills
by Oldowan hominins at FLK Zinjanthropus.
Passive scavenging by hominins from largely defleshed
felid kills could be falsified by two coinciding sets of observa-
tions on well-preserved bones typically defleshed later in the
consumption sequence (e.g., forequarters, neck; Blumen-
schine, 1986b). These bones should a) lack felid tooth marks,
with any tooth-marking having resulted from bone destruction
by hyenids or canids, and b) bear cut marks indicating bulk
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6 R.J. Blumenschine et al. / Journal of Human Evolution xx (2007) 1e7
defleshing by hominins. Before this test can be conducted, we
need to further explore the frequency and anatomical distribu-
tion of tooth-marking and butchery marking on carcasses
consumed to varying extents by carnivores. We also need to
be able to distinguish carnivore taxa using the tooth marks
and gnawing damage they produce (Haynes, 1983; Selvaggio
and Wilder, 2001; Domı́nguez-Rodrigo and Piqueras, 2003;
Pobiner and Blumenschine, 2003; Njau and Blumenschine,
2006). Finally, we need to distinguish cut marking associated
with bulk defleshing from that associated with removal of flesh
scraps (e.g., Domı́nguez-Rodrigo, 1997; Bunn, 2001; Egeland,
2003; Pobiner and Braun, 2005; Blumenschine and Pobiner,
2006).
Studies of bone surface modification on fossil assemblages
also need to adopt more rigorous procedural standards. As
a first step, blind testing of marks inflicted by known agents
on modern bone should become commonplace. Blumenschine
et al. (1996) remains the only published blind test of this na-
ture. Domı́nguez-Rodrigo did examine Blumenschine’s con-
trol collections, as he claims, but he did not submit to the
blind tests. Blind testing becomes especially important as po-
tential mimics for tooth marks and butchery marks become
known, but Domı́nguez-Rodrigo and Barba did not conduct
a blind test using specimens known to have been bioeroded
microbially or tooth-marked by carnivores. Blind testing
should also be expanded to include bones modified by insects,
because these marks are similar in size to carnivore tooth
marks, and some have a similar superficial morphology [see,
for example, Watson and Abbey, 1986; Kaiser and Katterwe
(2001); Fejfar and Kaiser (2005); Fig. 1b]. Finally, researchers
should demonstrate high inter-analyst correspondence in
surface mark identifications on fossil bones, such as Blumen-
schine and Capaldo have done for tooth marks and percussion
marks in their independent analyses of the FLK Zinjanthropus
long bones (Blumenschine et al., in press).
These efforts to improve the reliability and accuracy of sur-
face modification studies will avail verifiable inferences about
the nature of feeding interactions between hominins and other
members of fossil carnivore guilds, interactions that are central
to an understanding of hominins’ evolving anatomical, techni-
cal, and social capabilities (cf. Blumenschine and Peters,
1998; Blumenschine and Pobiner, 2006). A falsification of
passive scavenging from abandoned carnivore kills, such as
that purported to be shown by Domı́nguez-Rodrigo and Barba
(2006), requires the construction of robust neotaphonomic
models that are applied reliably to fossil assemblages. On
a number of fronts, Domı́nguez-Rodrigo and Barba (2006)
fall short of this methodological standard.
Acknowledgments
We are grateful to S.C. Antón, L.G. Blumenschine, J.A.
Cavallo, C.W. Marean, F. Mascia-Lees, B.L. Pobiner and three
reviewers, including T.R. Pickering, for comments on the tone
and content of this reply.
Please cite this article in press as: Robert J. Blumenschine et al., Carnivore tooth-marks, microbial bioerosion, and the invalidation of Domı́nguez-Rodrigo and
Barba’s (2006) test of Oldowan hominin scavenging behavior, J Hum Evol (2007), doi:10.1016/j.jhevol.2007.01.011
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