Journal of Archaeological Science 34 (2007) 1629e1640

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

Tooth marks and human consumption: ethnoarchaeological

mastication research among foragers of the Central African Republic
Matthew J. Landt*
Department of Anthropology, 150 College Hall, Washington State University, Pullman, WA 99164-4910, USA
Received 8 September 2006; received in revised form 2 December 2006; accepted 4 December 2006

Abstract

Taphonomically, much research has focused on the way in which predators and humans vie for calorically rich sources of protein. Anecdotal
evidence from ethnographies and experiments indicate that humans willdas do other obligate carnivores and omnivoresdmodify animal bones
with their teeth during consumption. Recent ethnographic research among the Bofi foragers of the Central African Republic provides an oppor-
tunity to explicitly understand the way in which humans imprint bone during mastication. This research identifies the signature of human tooth
marks on small mammal skeletons and addresses the way in which these marks may be archaeologically visible. The data presented herein sug-
gests that any model seeking to discuss the range of human dietary choices would be strengthened by considering the impact of humans in
zooarchaeological small fauna assemblages that may or may not have technological indicators of a human presence.
Ó 2006 Elsevier Ltd. All rights reserved.

Keywords: Taphonomy; Ethnoarchaeology; Mastication; Bofi foragers; Small mammals; Central African Republic

1. Introduction
To identify human involvementdand hence reliance upond
edible animal resources, archaeologists and paleoanthropolo-
gists rely on the visible impact of human activities on bone
remains during the acquisition and preparation of animal
protein for consumption (Dominguez-Rodrigo et al., 2005;
Egeland, 2003; Gifford-Gonzalez, 1989; Lupo, 1995; Lupo
and O’Connell, 2002; Lupo and Schmitt, 1997; Noe-Nygaard,
1989; Oliver, 1993; Potts and Shipman, 1981; Walker and
Long, 1977). By identifying the interactions of hominids in re-
lation to faunal remains, archaeologists are better prepared to
discuss the ways in which humans have come to interact with
the environment (Bird and Bliege Bird, 1997; Pulliam, 1981;
Sosis, 2002), acquire and compete for nutrients from carcasses
(Egeland et al., 2004; Pickering et al., 2004; Pobiner and
Blumenschine, 2003; Treves and Naughton-Treves, 1999), and
* Tel.: þ1 509 435 5080.
E-mail address: matthew_landt@wsu.edu
rely on each other (Boesch, 1994; Hawkes et al., 1997, 2001;
Hewlett, 1991; Knight, 2004; Marshall, 1994). Yet, regardless
of the cognitive mechanisms and adaptive benefits one ascribes
to early tool use and meat-sharing behaviors, it should be ex-
pected that hominids captured and consumed small animals
(aves, mammalia, reptilia, etc.) well before the point at which
they turned their attention to larger species (Bartholomew
and Birdsell, 1953; Fernandez-Jalvo et al., 1999; Goodall,
1963; Jones, 1984; Plummer and Stanford, 2000; Sept, 1992;
Winterhalder, 1997; Wynn, 2002; Yellen, 1991).
It is expected that small animals (<20 kg) are not processed
in identical ways to larger prey species (Dominguez-Rodrigo
and Barba, 2005; Tamplin et al., 1983; White, 1953;
Fancher and Landt, in preparation). The discontinuity between
small and large animal butchery techniques has been noted in
the literature, although the archaeological implications of such
have met with mixed results (Elkin and Mondini, 2001; Jones,
1984; Sept, 1992; Tamplin et al., 1983). Because small
animals can be butchered without the aid of tools, it may be
suspected that the bulk of small animal exploitation is not ar-
chaeologically visible at ephemeral occupation sites. Further,

0305-4403/$ - see front matter Ó 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2006.12.001
1630 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640
the vagaries of invasive rodent remains that do not show evi-
dence of human modification (i.e., burning, cut-marks, etc.)
continue to haunt archaeological interpretations (Speth,
2000; Stahl, 1982). Thus, numerous problems remain with
the archaeological identification of small animal comestibles
that do not show evidence of technological modification yet
where in fact exploited (cf. Madsen and Schmitt, 1998; Stahl,
1982).
Anecdotal evidence from numerous field studies amongst
the Dassanetch (Gifford-Gonzalez, 1989), Hadza (Oliver,
1993), Hottentot (Brain, 1981; Maguire et al., 1980), Nuna-
miut (Binford, 1978, 1881), and Sioux (White, 1953, 1955)
as well as two explicit exploratory studies that utilized collec-
tions of sheep in Argentina (Elkin and Mondini, 2001) and nu-
merous bird species in Hawaii (Weissler and Gargett, 1993)
indicate that humans will modify animal bones with their teeth
during consumption, as do other obligate carnivores and omni-
vores. However surficial the evidence may initially be for
archaeological application, it suggests a means of distinguish-
ing human and non-human accumulated small mammal assem-
blages. As Binford (1981:148) noted, ‘‘it is highly unlikely that
a normal hominid pattern of consumption would include gnaw-
ing to the extent that it would mimic the consequences of the
classic carnivore’s behavior.’’ As such, ‘‘there is no reason to
view, at least implicitly, the traces of gnawing as an exclusive
indicator of non-human animals’’ (Elkin and Mondini,
2001:260e261).
This research was undertaken to focus on the way in which
humans may leave archaeologically visible tooth marks from
consumption. By enveloping this project within a broader eth-
noarchaeological endeavor that focuses on the way in which
humans interact with small prey animals (Lupo and Schmitt,
1997, 2002, 2005) it is possible to further clarify the archaeo-
logical implications of human tooth marked small mammal
assemblages. The following essay provides descriptive and
contextual cultural and taphonomic information for human
tooth damage on the bones of four species of small mammals
(<20 kg) exploited by Bofi foragers in the winter of 1999e
2000 on the northern edge of the Congo Basin in the Central
African Republic. With realistic intent, this data is presented
in the hope of exploring ‘‘. how behavioral patterns that
are observed in individual cases can be understood as exam-
ples of the repertoire of hominid behavioral responses .’’
(Sept, 1992:22).
2. Background
The study site for the faunal assemblage used herein is near
the M’Baéré River at the village of Grima in the N’gotto Forest
Reserve in the southwestern portion of the Central African
Republic. The N’gotto Forest consists mainly of dense semi-
deciduous forest with pockets of naturally occurring open
wet-savanna and sections of raffia palm (Raphia sp.) swamp
forest located along the rivers. Because the N’gotto Forest lacks
extreme heat and cold fluctuations through the year,
reproduction for forest species is not limited to any specific sea-
son and the diversity of plants and animals in the forest remains
high. During the dry season in December 1999 and January
2000 roughly 150 Bofi foragers occupied a semi-permanent
village near Grima and/or a series of non-permanent hunting
camps established in the forest that were generally occupied
for a few days to a few weeks at a time (Lupo and Schmitt,
2002). The village of Grima was contemporaneously occupied
by approximately 200 Bofi farmers who continue to maintain
economic ties with the foragers based on exchanges of forest
products (e.g., koko leaves, payo nuts (Irvingia sp.), bush-
meat, etc.) and labor for wages and manioc (Fouts, 2002;
Lupo and Schmitt, 2002).
While the N’gotto Forest contains more than 115 species of
mammals (including 13 primate species), hunting by the Bofi
foragers is directed mostly towards 28 different mammalian
species of prey as well as a variety of birds, reptiles, fish, crus-
taceans, and insects (Dethier and Ghuirghi, submitted for
publication). Of the commonly pursued mammalian prey,
approximately 75% have a live weight under 20 kg, while
another three-quarters weigh less than 5 kg (Dethier and
Ghuirghi, submitted for publication; Hudson, 1990; Kingdon,
1974, 1982; Lupo and Schmitt, 2002, 2005; Noss, 1995). The
most common prey species, in the diet of the Bofi foragers are
blue duiker (Cephalophus monticola, 5 kg), brush-tailed por-
cupine (Atherurus africanus, 3 kg) and giant pouched rats
(Cricetomys gambianus, 1 kg) (Kingdon, 1974, 1982; Lupo
and Schmitt, 2002, 2005; Noss, 1995). Larger prey are pursued
when they are encountered, but these opportunities are rare.
Foragers who do not go into the forest for a foray may occa-
sionally hunt murid rats and mice (<1 kg live weight) in and
around the villages and camps.
The Bofi hunt year-round and employ a variety of individ-
ual and communal techniques, such as hand capture, snares,
traps, and nets to obtain meat. Most duiker meat is obtained
in communal net-hunts, a technique described in detail in pre-
vious literature (Lupo and Schmitt, 2002; Turnbull, 1965).
Porcupines, rats, and mice are more often captured by hand
while employing individual traps and/or fire drives. Details
of Bofi butchery practices are described in detail elsewhere
and need not be repeated here (Hudson, 1990; Lupo and
Schmitt, 2002; Noss, 1995; Zietz, 2002). The Bofi prepare
meat by either roasting the animal over an open fire, or by
boiling it in pots. The use of an open flame allows for easier
removal of hair and parasites and is mostly used with porcu-
pines, rats, and mice. The blue duiker is often boiled in
a pot with koko leaves or other vegetables. Duiker bones
may be chopped or hand fractured in order to ‘‘pot-size’’
them prior to boiling. These differential cooking patterns
may influence both the ability of the bone to record tooth
marks as well as the likelihood that individual foragers attempt
to acquire attached/within bone nutrients (Lupo and Schmitt,
1997; Speth, 2000). It is expected that the boiling of duiker
bones may soften the cortical layers and increase the rate of
tooth mark recordation at the same time that it will expedite
muscle removal and decrease the likelihood of mastication
events being recorded. Conversely, it is expected that the roast-
ing of the porcupine, rat and mice carcasses may dry and
strengthen muscle attachments, thus increasing the relative
 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640
likelihood of teeth imprinting on a bone’s surface. Hence, dif-
ferential meal preparation is recognized as a potential factor
for distinguishing variation in the Bofi faunal assemblage.
Information on daily Bofi forager hunting and subsistence
activities was gathered through observations (via focal fol-
low), interviews, and bones collected from individual meals.
Daily systematic assemblages of bone refuse were collected
directly from the consumer and/or the consumer’s family by
project members who exchanged empty bags for bags with
the remains of meals. These faunal collections were made
both in the semi-sedentary forager camp near Grima and in re-
mote camps. The collected assemblage consists of blue duiker,
giant pouched rat, brush-tailed porcupine, murid mice, civet
(Civettictis civetta), land tortoise (Kinixys erosa), and pangolin
(Phataginus tricuspis). By collecting bones directly from the
consumer, bones were never exposed to forager dogs and
thus avoided the complicating influence of canid attrition
and destruction (Binford, 1981; Hudson, 1990; Munson and
Garniewicz, 2003; Payne et al., 1985; White, 1992; Zietz,
2002). The collected bone specimens were then cleaned by
hand, disinfected, dried, examined, and recorded in the field
before being transported to the zooarchaeological laboratory
at Washington State University. During field cleaning, re-
searchers utilized locally available pot scrubbers as mechani-
cal abrasives in removing any remaining soft tissue before
drying and transportation. As such, it is recognized that field
cleaning may have impacted this collection of human tooth
damaged faunal remains.
The author undertook an exploratory study with human
gnawed rabbit remains to clarify the impact of mechanical
abrasives on tooth mark damage. The details of the study
are made explicit elsewhere (Landt, 2004), but resulted in
visible polishing and scoring. The scoring associated with
cleaning produced small isolated or grouped linear scratches
that appear as elongate v- or u-shaped grooves with smooth
internal surfaces. Most of these striations are not visible to
the unaided eye and are only noticeable at microscopic
levels (cf. Shipman and Rose, 1983). Measurements on
cleaning scratches in the study group indicate that any rel-
atively small linear groove (ranging from 0.01 to 0.10 mm
in width), whether it is isolated or in a cluster (parallel,
perpendicular or angled to the majority of marks) in the
Bofi faunal assemblage, may have been produced by post-
collection scrubbing.
Since individual human tooth mark identifications are
made based on a combination of macro- and microscopic
identification, it is possible to examine each bone for evi-
dence of field cleaning while examining them for evidence
of mastication damage. Thus, tooth marks that are visibly as-
sociated with field cleaning can be identified and their size
can be contrasted against those marks that are not associated
with field cleaning. Discussion regarding any variation in
tooth mark size caused by field cleaning is further detailed
in the text.
In this study of human mastication remains it is important
to note that some Bofi foragers practice traditional tooth shap-
ing of the maxillary incisors when they enter adulthood.
1631
However, the Bofi foragers do not practice shaping of their
mandibular incisors as do their Aka neighbors (Walker and
Hewlett, 1990). To fully shape the maxillary incisors, the for-
agers chip or break off both the mesial and distal corners,
leaving the incisor pointed from the occlusial surface to the
gum-line. Field observations indicate that there is a great
deal of variability in tooth shaping practice amongst the for-
agers and that not all shaped incisors are ‘fully’ shaped. In
a survey of 54 adolescent and older (216 potential upper inci-
sors) Bofi foragers near the village of Grima from October to
November 2003, three-quarters (n ¼ 42) of the population had
one or more modified upper incisor(s) (Landt, 2004). Anec-
dotal observations indicate that when humans chew on bone
(whether to gnaw on the bone itself, or in the process of re-
moving tough adhering tissue), it is the premolars and molars
that are most often used and not the incisors, thus inherently
recognizing that the first molars are the point of peak mastica-
tory muscle force (Kieser, 1999). However, it is possible that
the Bofi forager practice of tooth shaping may impact the fau-
nal assemblage used herein in some unforeseen or incidental
way as the relative distance between teeth and tooth rows is
quite small in humans as compared to other carnivores and
broken tooth edges may result in microscopic morphological
variations (i.e., internal striations). Additional studies of
human mastication amongst peoples with unshaped teeth
would provide appropriate comparative documentation. Until
such studies occur, it is important to reiterate that ‘‘like all
lives, they can be used as examples or serve as representative
types. But ultimately they are unique, individual, [and] impos-
sible to define or replace .’’ (Schlosser, 2002).
3. Faunal assemblage and methods
Taken as an aggregated assemblage, the Bofi faunal col-
lection exhibits an array of damage types from field butcher-
ing, culinary processing, and consumption (i.e., cut marks,
chop marks, tooth marks, and fracturing). While a singular
focus on one damage type in an archaeological assemblage
is a poor methodology for understanding the complexity of
taphonomic influences, the point here is to isolate and de-
scribe a previously under-represented type of human process-
ing damage. Thus, the differential distribution of multiple
damage types (i.e. cutmarks, burning, etc.) across the assem-
blage, and their localized relationship to one another is de-
scribed and analyzed elsewhere (Fancher and Landt, in
preparation).
For human tooth marks, assignment of ‘tooth marked’ and
‘non-tooth marked’ were based on micro- and macroscopic
criteria established by Binford (1978, 1981), Blumenschine
and Selvaggio (1988), Bonnichsen and Will (1990), Brain
(1981), Capaldo and Blumenschine (1994), Fisher (1995),
Haynes (1980, 1983), Johnson (1989), Pickering and Wallis
(1997) and most recently Njau and Blumenschine (2006)
with special attention given to the descriptions of Elkin and
Mondini (2001) and Weisler and Gargett (1993). The charac-
teristic definitions used herein are made explicit in Table 1
with comments on typical skeletal element associations. Using
1632 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640
Table 1
Definitions and characteristics used in identifying tooth marked bones
Type(s) of damage Description(s)
Crenulated edges e Identified by broken edges that bisect all bone
layers and exhibit internal crushing along the broken
margins. Often associated with indicators of localized
forces (i.e., pits, punctures, notches, scoring)
e Commonly associated with bones that have relatively
thin cortical bone layers overlying cancellous matrices
(e.g. rib shafts, innominates, and vertebral processes)
Fractured edges e Identified by broken bone edges that bisect all bone
layers, do not exhibit internal crushing of the
margins as do crenulated edges, and are often
unassociated with other types of damage
e Associated with all bones although ribs and
appendicular elements tend to be fractured as a result
of ‘‘pot-sizing’’ for consumption
Tooth pits e Indentation on the surface of the bone with visible
crushing of cortical bone layer(s) along the margins of
the impression. Does not breach the entire cortical
surface, leaving the majority of the outer margin of the
impression intact
e Typically shallow in cross-section and ovoid in plan
view though they are frequently found in irregular
shapes
e Commonly associated with relatively thick cortical
bone layers (e.g., shaft portions of appendicular
elements)
Tooth punctures e Characteristically results in an indentation that
breaches the entire cortical bone surface yet leaves
the outer surface margins intact (although radial cracks
may be visible). Visible crushing of cortical and
cancellous bone layer(s) often exists along the interior
margins of the impression
e Mostly round to ovoid in plan view
e Typically associated with thin cortical bone layers
that overly cancellous matrices (e.g., ribs, vertebrate,
distal and proximal portions of appendicular elements)
Notches e Identified as an indentation that breaches the cortical
bone surface due to compression forces and results in
a bisected margin. Visible crushing of cortical and
cancellous bone layer(s) may exist along the upper
intact margins of the impression
e Associated with relatively thick cortical bone layers
(e.g., medial shafts of appendicular elements)
Scoring e Identified by elongated impressions in the cortical
matrix. The impression is typified by a rounded
cross-section that occasionally shows internal crushing
along the borders. Lacks parallel striations associated
with cut and trampling marks
e Often associated with other damage types
Each description entails the characteristics of identification and typical skeletal
element associations.
these definitions as guidelines for mastication damage from
the Bofi foragers, roughly one-fifth of the 2514 bones in the
1999e2000 Bofi faunal assemblage are tooth marked
(n ¼ 452, Table 2).
A random sample of 111 tooth marked bones (24.5%),
stratified by aggregated cultural butchery divisions following
skeletal elements for each species (i.e., forelimb bones, hind-
limb bones, axial halves, head and neck elements; see
Hudson, 1990; Landt, 2004 for details), was used for SEM
analysis in an attempt to represent differential tooth mark
patterning that may exist between skeletal elements because
of bone density or cultural habits (cf. Selvaggio and Wilder,
2001), and to gain an accurate picture of the size and skeletal
patterning of human tooth marks. The bones selected as part of
this sample were initially viewed with a 10 hand lens and
strong lighting to identify areas with and without modification.
Analysis of those areas was then supplemented with an image
analysis workstation (IAW) and a scanning electron micro-
scope (SEM). Both the IAW and SEM are housed in the Elec-
tron Microscopy Center, Department of Biological Sciences at
Washington State University, Pullman, WA. The IAW consists
of a Wild-Heerburg Dissecting Scope that is connected to
a Color Image Analysis CCD MicroImage videosystem where
the elusive image is collected with an NIH Image capturing
system. Preparation of the bones for microscopic analysis con-
sisted only of dehydration and desiccation to remove excess
water and grease to meet the vacuum requirements of the
SEM specimen chamber as skeletal elements from small mam-
mals need not undergo the additional trials and tribulations of
large specimen SEM work (Gilbert and Richards, 2000). Mea-
surements of tooth mark damage for pits and punctures were
taken from the SEM photos as the maximum linear dimension
(MLD) of each mark. The maximum breadth of each tooth pit
was also measured, but as the ranges covaried with the MLD
(cf. Dominguez-Rodrigo and Piqueras, 2003), only the MLD is
reported here. Scoring marks were measured as the maximum
breadth of the scratch and tooth notches were measured fol-
lowing Capaldo and Blumenschine (1994).
4. Damage descriptions
The most obvious types of macroscopic damage on the
bones resulting from human mastication activities are crenu-
lated edges (Fig. 1), tooth punctures, tooth pits (Fig. 2), and
tooth scoring (Table 3). Crenulated edges are often associated
with other carnivore damage patterns (e.g., pits, punctures,
scoring, etc.) and exhibit crushing along the interior of broken
margins. Broken edges that do not exhibit interior crushing of
the margin and are not associated with other damage types are
classified as fractured edges and are typically associated with
hand fracturing related to the ‘pot-sizing’ of skeletal elements.
Tooth pits are shallow depressions that do not completely
breach cortical bone layers. They were recorded with a variety
of plan views from circular to irregular, though all had crush-
ing evident around the margins. Some tooth pits and scoring
are macroscopically visible, yet many pits and scores were
only visible microscopically (Fig. 3). Tooth scoring (Fig. 4)
is identified by elongated impressions in the cortical bone sur-
face. These elongate impressions tend to have u-shaped cross-
sections without internal crushing though some may be visible
along the margins. Tooth scoring is often associated with tooth
pits and crenulated edges. Tooth punctures were defined by
a complete breaching of cortical bone layers that leaves the
bone surface outside of the depression intact. As with tooth
pits, punctures leave visibly crushed bone layers along interior
margins. Tooth notches are defined by the complete breaking
of cortical bone layers that result in a bisecting of the bone
 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640 1633

Table 2
Raw NISP counts of tooth damaged bones in the 1999/2000 Bofi forager faunal assemblage by species used in this analysis with associated percentages
Element group Blue duiker Pouched rat Brush-tailed porcupine Murid mice Totals
NISP % NISP % NISP % NISP % NISP %
Head and necka 26 10.24 5 3.91 0 0.00 1 3.70 32 7.08
Post-cervical axiala 155 61.02 70 54.68 46 63.01 7 25.93 278 61.50
Forelimba 33 12.99 35 27.34 20 27.40 2 7.41 90 19.91
Hindlimba 14 5.51 18 14.06 7 9.59 13 48.15 52 11.50
Total tooth marked 228 19.84 128 18.31 73 25.17 23 6.12 452 17.98
Total non-tooth marked 921 80.16 571 81.68 217 74.83 353 93.88 2062 82.02
Total assemblage 1149 699 290 376 2514
a
Percentages of the element groups are proportions of the tooth marked assemblage only.

layers at the point of impact. Tooth notches are often associ-
ated with diaphysis portions of appendicular elements and as
such do not often record crushing along interior margins
though some crushing may be apparent at the upper interior
portion of the point of impact.
Tooth pits, tooth punctures, and tooth notches are all per-
petuated by a force that is driven perpendicular to the surface
of the bone. As such, differences in their characteristic defi-
nitions rely in part on the varying composition of the skeletal
element and its ability to record tooth impressions as well as
the relative amount and differences in force. Since tooth pits
rely on a relatively minimal amount of force, they can be lo-
cated along any portion of individual skeletal elements.
Tooth punctures, by the nature of the damage, must be found
on skeletal elements, and portions thereof, with relatively
thin cortical bone layers underlain by cancellous bone matri-
ces (e.g., ribs and epiphyseal ends of appendicular elements).
Tooth notches are associated with diaphysis fragments as
thick cortical bone layers are more responsive to the prereq-
uisite compression forces. As the same tooth can create any
and all of the above tooth marks, any distinctions within
types of tooth damage (i.e., differences between tooth pits)
are more a product of the interacting components (e.g., dif-
ferent enamel and bone matrices) than they are the individual
actors per se. This point is made clear by Njau and Blumen-
schine (2006:149) with their recognition of bisected tooth
marks and a specific reptilian tooth structure. The marks re-
corded here for the Bofi foragers, with their omnivorous
tooth structure, do not differ markedly from other mamma-
lian carnivores in the strict morphological sense of impres-
sions left on bones with teeth.
While morphological differences are unlikely to exist be-
tween predators with similar tooth structures on prey of similar
sizes, it is not unreasonable to expect that the distribution, fre-
quency, and size of tooth mark damage across varying prey
size classes may yet clarify the size range of actors if not
the actor itself (Dominguez-Rodrigo and Piqueras, 2003;
Njau and Blumenschine, 2006; Pickering and Wallis, 1997;
Pickering et al., 2004; Selvaggio and Wilder, 2001). With
that in mind, the following descriptive account provides qual-
itative and quantitative information regarding one data point
within a range of variation for human tooth mark damage on
four species of small mammals.

5. Distribution

Qualitatively, the blue duiker bones with human mastication

Fig. 1. SEM image of crushing along a broken margin of the ventral process of
a pouched rat vertebrate at a magnification of 20.
damage are largely complete. The majority of mastication
damage on blue duiker long bones focused on the removal of
minimal to moderate amounts of cancellous bone tissue (e.g.,
removal of trochanters, and ends of ribs). Following divisions
of appendicular long bones by Bunn (2001), tooth mark dam-
age on blue duiker bones is largely localized to the proximal
and distal ends, where only one of the appendicular long bones
in the sample (n ¼ 36) recorded tooth mark damage to the di-
aphysis portion. On irregular or flat duiker bones, tooth mark-
ing is focused on the edges of the bones. The crenulated edges
of mastication damaged pouched rat remains is similar to that
described by Weisler and Gargett (1993) where entire long
bone epiphyses are removed as well as the ends of other bones
(e.g., innominate crests, rib halves). Two of the giant pouched
rat appendicular long bones (n ¼ 44) recorded tooth markings
1634 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640
Fig. 2. SEM image of tooth pits by a Bofi forager on the innominate of a giant
pouched rat at a magnification of 20 (A) and 50 (B).
(pits and scorings) on the medial shaft. The porcupine bones
damaged during mastication are mostly complete and damage
to appendicular long bones is entirely limited to the proximal
and distal ends where small to moderate amounts of cancellous
bone tissue were removed. As with the other species, tooth
marking on the small mouse bones is limited to the proximal
and distal ends of long bones. This damage is discernible in
crenulated edges that resulted in the removal of small amounts
of cancellous bone tissue (i.e., the femoral greater trochanter)
but did not result in the removal of epiphyseal ends.
6. Frequency
The post-cervical axial bones in the Bofi faunal assemblage
are the most frequently gnawed elements in all but the mice
remains. The apparent focus on axial elements (e.g., ribs and
vertebrate) amongst the larger animals is likely linked to the
relatively high number of muscle attachments and muscle
groups compared to appendicular elements. The Bofi tend to
significantly gnaw on more axial blue duiker elements than
other portions of either the giant pouched rat or brush-tailed
porcupine (c2 ¼ 29.36, df ¼ 6, p  0.001). In fact, 43% of
the chi-square value is provided for by higher than expected
values of tooth marking on blue duiker axial elements and
lower than expected values on appendicular elements. The in-
creased focus on axial bones among the blue duiker remains
(especially the cervical vertebrate) is likely a product of shar-
ing and redistribution. Blue duiker are often captured in com-
munal net hunts and divided amongst participants whereas the
capturing of pouched rat, porcupine and mice are less likely to
be a communal activity (Lupo and Schmitt, 2005). The head
and attached cervical vertebrate of a net caught blue duiker
are often given as a unit to the person who first seizes the
prey (Hudson, 1990). As an individual unit of distribution
and consumption the blue duiker head and neck region may re-
ceive more mastication focus than do the head and neck of
prey species which are not similarly divided and redistributed.
This is further supported by the statistically similar distribu-
tional frequencies (c2 ¼ 4.16, df ¼ 3, p  1) of giant pouched
rat and brush tailed porcupine. A number of future research
controls that account for differential meal preparation tech-
niques and redistribution networks would strengthen the ideas
suggested here.
The relatively low percentage of tooth marked mouse
bones in the assemblage is also a product of both the method
of consumption and the physiological nature of mouse bones.
Mice are typically roasted over an open fire and the hair is
scrapped off before an individual tears portions free and con-
sumes the meat. The Bofi tend to choose the hindlimb por-
tion for consumption as noted by the significantly high
number of hindlimb mouse elements with gnawing damage
(c2 ¼ 77.04, df ¼ 9, p  0.001) that is associated with the
relatively large quadricep and gluteal muscles. During con-
sumption, a relatively small amount of pressure is needed
to fracture mouse bones, such that mouse bones are more
likely to fully collapse under pressure from the compara-
tively large Bofi teeth than to remain intact and record tooth
impressions (cf. Selvaggio and Wilder, 2001). Further, as
mice are not butchered with any tools, many bones, espe-
cially tarsals and carpals, remain articulated and uneaten,
and were recovered during the ethnographic interviews.
The collection of uneaten mouse elements bolsters the total
numbers of mice elements in the collection and reduces the
overall percentage of tooth marked bones.
 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640 1635

Table 3
Mastication damage types within the Bofi forager faunal SEM sample
SEM sample Crenulated Fractured Pitsb Puncturesb Notchesb Scoringb Bones with
edgesa edgesa evident
field cleaning
Blue duiker (n ¼ 55) 88 14 38 6 e 38 33
Giant pouched rat (n ¼ 26) 37 13 23 e 8 18 16
Brush-tailed porcupine (n ¼ 21) 32 3 16 1 5 17 16
Murid rats and mice (n ¼ 9) 5 6 3 e e 4 4
Total counts (n ¼ 111) 162 36 80 7 13 91 71
a
Reported crenulated and fractured edges in numbers of individual occurrences.
b
Reported damage is recorded as presence/absence of bones with damage type.

7. Tooth mark measurements
The range of tooth pit MLDs are recorded in Table 4. The
smallest measured tooth pit in the assemblage was measured
on a blue duiker rib and the largest measured tooth pit was on
a blue duiker tibia. An independent sample T-test using
SPSSÒ indicates that on blue duiker bones, the range of varia-
tion for the MLD of tooth pits associated with clear evidence
of field cleaning is significantly smaller than those that are not
(t ¼ 3.432, df ¼ 93, 2a ¼ 0.001), while pits on the pouched
rat and brush-tailed porcupine are not significantly different be-
tween those with and without evidence of cleaning (t ¼ 0.448,
df ¼ 68, 2a ¼ 0.656 and t ¼ 0.619, df ¼ 33, 2a ¼ 0.540 re-
spectively). Although field cleaning altered the surfaces of
bone in the Bofi assemblage, it does not appear to have substan-
tially altered the overall morphology and hence identification of
tooth mark damage.
Removing tooth pits associated with the well-cleaned
blue duiker bones from further analysis, T-tests indicate that
the range of tooth pit sizes is significantly different between
those on blue duiker and on pouched rat remains (t ¼ 3.160,
df ¼ 110, 2a ¼ 0.002), brush-tailed porcupine remains (t ¼
3.026, df ¼ 75, 2a ¼ 0.003), and the murid mice remains
(t ¼ 1.751, df ¼ 45, a < 0.05). A comparison between tooth
pits sizes on pouched rat and brush-tailed porcupine remains
indicates that the ranges are not significantly different (t ¼
1.024, df ¼ 103, 2a ¼ 0.308). However, the size ranges are
significantly different on the pouched rat and the murid mice
remains (t ¼ 1.641, df ¼ 73, 2a ¼ 0.105) as well as between
tooth pits on the murid mice and brush-tailed porcupine re-
mains (t ¼ 1.785, df ¼ 38, a < 0.05). Thus, even though the
average tooth pit size across the assemblage varies less than
a single millimeter, inter-species variations may themselves
be distinct.
The significant differences between tooth pit size ranges
across four different sized species of animals (blue dui-
ker ¼ 5 kg, brush-tailed porcupine ¼ 3 kg, pouched rat ¼ 1 kg,
and the murid mice < 1 kg) presented here suggests that
dimensions of tooth pits on small mammal remains are partly
dependent on the size of the bone being gnawed (cf. Selvaggio

Fig. 3. SEM image of irregular tooth pits on the pubic ramus of a giant
pouched rat. Fig. 4. SEM image of a tooth scratch with a tooth pit on a blue duiker tibia.
1636 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640

Table 4 Table 5
Size ranges of tooth pits measured as maximum linear dimension in Size measurements of tooth scratches as maximum breadth in millimeters
millimeters n Minimum Median/mean Maximum Std. dev.
n Minimum Median/mean Maximum Std. dev. Duiker a
42 0.053 0.218/0.268 0.630 0.127
Duikera 42 0.205 0.909/1.437 4.310 1.220 Duikerb 36 0.017 0.086/0.086 0.147 0.039
Duikerb 53 0.135 0.621/0.803 2.480 0.507 Rata 22 0.050 0.132/0.152 0.343 0.087
Rata 22 0.179 0.721/0.853 1.990 0.520 Ratb 76 0.004 0.040/0.042 0.121 0.030
Ratb 48 0.193 0.757/0.920 3.214 0.604 Porcupinea 13 0.027 0.143/0.168 0.466 0.115
Porcupinea 5 0.332 0.714/0.692 1.327 0.398 Porcupineb 56 0.005 0.026/0.031 0.086 0.020
Porcupineb 30 0.227 0.744/0.805 1.671 0.374 Micea e e e e e
Micea 4 0.200 0.393/0.529 1.130 0.436 Miceb 17 0.005 0.027/0.030 0.086 0.026
Miceb 1 e 0.223 e e Totala 77 0.027 0.186/0.218 0.630 0.126
Total 205 0.135 0.750/0.955 4.310 0.754 Totalb 185 0.004 0.036/0.046 0.147 0.035
a
Measurements of tooth pits where bone had no visible evidence of field Total 262 0.004 0.057/0.097 0.630 0.108
cleaning. a
b Measurements of tooth pits where bone had no visible evidence of field
Measurements of tooth pits where bone surface was clearly marked by
cleaning.
field cleaning. b
Measurements of tooth pits where bone surface was clearly marked by
field cleaning.

and Wilder, 2001), and hence of the animal being consumed.

Given that tooth mark sizes are partially limited by the size of generally occur as single or paired marks near crenulated
the skeletal elements for small animals, this research indicates edges, tooth punctures and tooth pits.
that humans and other carnivores are likely to leave similarly Previous reports have indicated a mean tooth score breadth
sized marks on small prey remains. This is not surprising as on diaphyseal skeletal elements between 0.2 and 2.2 for a vari-
one might expect the interaction of enamel with bone to perpet- ety of carnivores (Dominguez-Rodrigo and Piqueras, 2003;
uate a similar reaction once the size, strength, and density of Njau and Blumenschine, 2006). The average Bofi forager
both the bone and the actor are controlled (Binford, 1981; tooth score breadth (0.03e0.63 mm) falls at the very lower
White, 1992). range of these means and is most similar to tooth scores left
Dominguez-Rodrigo and Piqueras (2003:Table 1) reported by jackals on small-sized carcasses where the 95% confidence
a series of diaphyseal tooth pit measurements for a variety of interval is 0.19e0.73 mm (Dominguez-Rodrigo and Piqueras,
carnivores where the mean length is ca. 1.5e4.0 mm and the 2003:Table 2). However, tooth scores left by hyaenas, ba-
overall weighted mean is 2.67 mm. The size range for Bofi for- boons, bears and German Shepherds on larger prey remains
ager tooth pits (0.14e4.31 mm) clearly coincides with the all had tooth score marks that fell within the range described
overall tooth pit range of jackals where Dominguez-Rodrigo here for the Bofi (Dominguez-Rodrigo and Piqueras, 2003).
and Piqueras (2003) report a 95% confidence interval of Table 6 provides the overall size ranges of tooth punctures
0.51e4.2 mm when the diaphyseal and epiphyseal ranges are and tooth notch breadth and total notch ratio in this collection.
combined. The overlapping size of tooth pits between the The largest puncture was on the proximal end of a blue duiker
Bofi foragers and jackals is even more remarkable as the jackal tibia and the smallest puncture mark was left on a blue duiker
was the only carnivore in the case study to have feed on a small
sized carcass (Dominguez-Rodrigo and Piqueras, 2003:1386).
Further, the rest of the carnivores all left tooth pits smaller Table 6
than 4.0 mm in length on medium-sized bovid and equid re- Size ranges of tooth punctures measured as maximum linear dimension in mil-
limeters and tooth notch breadth dimensions
mains (Dominguez-Rodrigo and Piqueras, 2003).
The range of maximum breadth measurements for tooth n Minimum Median/mean Maximum Std. dev.
scratches is provided in Table 5. Two hundred sixty-two Puncture
tooth scratches were measured with the largest being less Duikera 7 0.874 1.018/1.949 4.180 1.324
Duikerb 9 0.291 1.360/2.191 4.310 1.673
than three-quarters of a millimeter in width and the narrowest Porcupinea e e e e e
measuring less than one-tenth of a millimeter wide. Given Porcupineb 1 e 2.600 e e
the results of the pilot study noted earlier, many of these Total 17 0.291 1.98/2.232 4.310 1.444
measurements are impacted by or derived from the field Notch Breadth
cleaning methodology. Excepting those scratches that are as- Rata 6 0.714 1.516/1.888 4.600 1.447
sociated with field cleaning, the mean tooth scoring mark Ratb 4 0.482 1.478/1.359 2.000 0.654
centers at approximately one-twentieth of a millimeter in Porcupinea 1 e 3.670 e e
Porcupineb 4 0.584 1.393/2.130 5.150 2.115
width (n ¼ 77) while the overall range on these small prey
Total 15 0.482 1.670/1.930 5.150 1.456
animals does not exceed three-quarters of a millimeter. The Notch ratio 15 0.734 1.977/2.238 5.255 1.233
majority of tooth scratches measured here, whether associ- a
Measurements of tooth pits where bone had no visible evidence of field
ated with field cleaning practices or not, are imperceptible cleaning.
without the aid of magnification. The scratch marks that do b
Measurements of tooth pits where bone surface was clearly marked by
not appear associated with damage from field cleaning field cleaning.
 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640
thoracic vertebrate. The lone puncture on the remains of
brush-tailed porcupine was located on the iliac portion of an
innominate. Intra-species comparisons of the range of varia-
tion in tooth punctures on blue duiker elements that evidence
high amounts of cleaning damage are not significant in this
case (t ¼ 0.314, df ¼ 14, 2a ¼ 0.758). Again, the range of
variation for Bofi forager tooth punctures most closely resem-
bles that reported for jackal gnawed remains where the epiph-
yseal pits range from 2.8 to 4.2 mm and hyaenas, baboons,
bears, and dogs all leave epiphyseal pits under 4.0 mm in
size (Dominguez-Rodrigo and Piqueras, 2003).
Tooth notches ranged in breadth from less than 0.5 mm on
a pouched rat thoracic vertebrate to over 5 mm on a brush-
tailed porcupine fibula. Although tooth notches display a large
size range in breadth, the ratio of notch breadth to depth de-
fines them as more semicircular than arcuate (Capaldo and
Blumenschine, 1994). Thus, while distinctly smaller than
any of Capaldo and Blumenschine’s sample of tooth notches
on bovid remains, the semicircular nature of the notches places
them well within the range of carnivore behavior (Capaldo and
Blumenschine, 1994:Table 4).
8. Discussion
Surprisingly, the largest tooth marks in the Bofi forager fau-
nal assemblage are not on the bones of the heaviest animal.
The blue duiker (5 kg) remains exhibit tooth pits around
4 mm in maximum size, yet the pouched rat (1 kg) and porcu-
pine (3 kg) bones both exhibit tooth notches that are around
5 mm in breadth. The difference in tooth marks may be due
to the overall robust morphology and size of the individual an-
imal elements, where the shorter but relatively broader rat and
porcupine bones are more apt to retain larger tooth marks as-
sociated with full breaching of bone walls, than are the lithe
but longer blue duiker elements. Similarly, the relatively frag-
ile and noticeably smaller mouse bones (<1 kg live weight)
are unlikely to retain tooth marks of any comparable size, as
in this assemblage where only a few small tooth pits and tooth
scores were recorded.
While the physical difference between the average size of
tooth pits on the murid mice and the blue duiker remains is
less than a single millimeter, these differences are expected
to be exacerbated when medium to large prey animals are in-
cluded in the sample. This is not meant to suggest that human
gnawing on giraffe elements will be confused with hyaena
gnawing on the same elements, as the maximum size of tooth
marks is likely a proxy that can be used to eliminate smaller
carnivores from a list of potential actors on large prey remains.
However, given the overlap in tooth mark size ranges between
the Bofi and a series of other larger carnivores, it does suggest
that on small mammal remains, the maximum tooth mark size
is limited more by the ability of individual bone elements to
record tooth impressions, than by the size of the actors tooth.
Dominguez-Rodrigo’s and Piqueras’ (2003) study does
suggest that the mean size of tooth pits on epiphyses and di-
aphyses may be useful in distinguishing coarse size classes
of consumers though it is unlikely to indicate a specific
1637
predator (Elkin and Mondini, 2001; Lyman, 1994). If such is
the case, tooth pit sizes in the faunal collection analyzed
here would indicate that the Bofi leave tooth marks that are
similar to small and medium-sized carnivores (i.e., small
canids, medium felids and baboons) (Dominguez-Rodrigo
and Piqueras, 2003). However, since the maximum size of
a given tooth mark is controlled by (1) the maximum size of
the predators tooth, (2) the overall size and robusticity of the
damaged skeletal element, as well as (3) micro-morphological
differences in structural composition and bone compression
(e.g., cancelleous ribs of bovids versus cortical long bones
of rodents), the overlap of tooth pit dimensions across multiple
carnivorous species is considerable in the small tooth mark
size category (cf. Selvaggio and Wilder, 2001; Dominguez-
Rodrigo and Piqueras, 2003). More simply, if the robusticity
and overall size of the bone element is in part responsible
for the size of the tooth mark, then tooth pit sizes across small
mammal species (<20 kg) is unlikely to be indicative of a size
class of consumer let alone of a specific predator (cf.
Dominguez-Rodrigo and Piqueras, 2003; Shipman, 1983).
Thus, not only are human tooth marks on small animals likely
to be confused with those of small canids on similarly sized
remains, but the size of the mastication damaged prey is an
important variable that needs to be controlled in future
research.
Intra-assemblage patterning in the Bofi faunal assemblage
indicates that small sized prey receive damage from human
consumption across the entire skeleton (Table 2), though the ax-
ial elements receive more focus. Further, minimal mastication
damage focusing on the ends of long bones is not likely to be
focused on attaining access to grease or marrow from within
the bone in the case of the boiled blue duiker. The focus of mas-
tication damage on long bones in the Bofi assemblage appears
to emphasize the removal of attached tissues as evidenced by
the high percentage of mastication damage on points of muscle
attachment in concert with the minimal destruction of the bone
(i.e. the consumptive removal of only the greater trochanter on
12 of 13 mouse femurs that is intuitively associated with the
gluteal abductor muscle attachment). There are few reports
that focus on features of carnivore tooth mark patterning
across small (<20 kg) mammal carcasses (for exceptions see
Andrews, 1990; Andrews and Nesbit Evans, 1983; Fernandez-
Jalvo and Andrews, 1992). Understandably, this is because
a large number of predators (canids, felids, mustelids, viverrids)
consume entire carcasses of small mammals (Andrews, 1990;
Hernandez et al., 2002; Kruuk, 1972; Schmitt and Juell,
1994). Thus, incidental tooth damage on points of muscle
attachments across small mammal skeletons, which are not
deposited as parts of fecal assemblages, may be understood as
part of a range of human consumptive behavior patterns.
Proof of this hypothesis will be archaeologically attainable
when ethnographic, experimental and archaeological assem-
blages identify bones damaged during consumption events
with information regarding the size of the prey remains, the
overall amount of bone destruction (e.g., mostly complete ver-
sus highly fragmented), the frequency of mastication across
the collection (e.g., 6% of gnawed mice remains, 25% of
1638 M.J. Landt / Journal of Archaeological Science 34 (2007) 1629e1640
gnawed porcupine remains) and the skeletal patterning of that
gnawing (e.g., evenly across the skeleton or focused on spe-
cific elements) as indicators of which predator is likely to
have left evidence of their consumption activity. In this sense,
it may be surprising to find a tooth damaged small mammal
bone collection like that seen for the Bofi (i.e., tooth marking
on roughly 20% of the assemblage with a non-destructive fo-
cus on axial elements and long bone epiphyseal portions) in
association with other predators that are not etched by diges-
tive acids and highly fragmented. Thus, while Bofi consump-
tion damage appears to focus more on soft tissue than on
within bone nutrients, the frequency and intensity of gnawing
in relation to the size of the skeletal element and the overall
assemblage may well prove to be a human consumptive signa-
ture in small mammal faunal assemblages. While not suggest-
ing that humans cannot consume bone, this research indicates
that modern humans may be archaeologically distinctive from
other carnivores in that the full consumption of large or small
animal bones is not a typical part of modern human nutrition
acquisition.
9. Conclusions
The samples analyzed in this paper are useful in examining
the morphological indication of human subsistence activities
and the way those faunal remains may or may not differ
from the mastication events of other obligate carnivores and
omnivores. This research suggests that the size of tooth
mark damage produced by human mastication on the surface
of small mammal skeletal elements is characteristically iden-
tical in size and morphology to damage produced by other
predators (cf. Elkin and Mondini, 2001). This appears to be
a product of the nature of the interacting components and
was foreshadowed by Binford (1981) and White (1992:155)
who noted that, ‘‘It is evident from simple mechanical consid-
erations that substantial overlap between human and carnivore
chewing damage on bones will be shown by future research in
this area.’’ In other words, there are a limited number of ways
in which tooth enamel, regardless of size, can interact with
bone matrices. Thus, as seen via the Bofi forager faunal collec-
tion, consumption of small animal remains by carnivores and
hominids is likely to produce the same archaeological signa-
ture (i.e. crushed margins, pits, punctures, etc.), although the
patterning of that damage is likely to be different. It may be
better understood by noting that a carnivore (be it a hyaena
or weasel) in the throes of sensitivity could produce the
same minimalistic damage on mouse bones that we see in
the Bofi forager faunal collection. The fact that hyaenids, ur-
sids, felids, canids, mustelids, and other carnivores do not gen-
erally produce this type of damage and will often consume
entire carcass of small mammals (Andrews, 1990; Andrews
and Nesbit Evans, 1983; Hudson, 1990; Lyon, 1970; Payne
et al., 1985; Willey and Snyder, 1989) is an interesting point
worthy of further research attention.
This work was initially undertaken in an effort to expand
upon the multitude of ways in which archaeologists identify
cultural patterning in archaeological assemblages and increase
the visibility of human involvement in faunal assemblages. As
this paper represents only one usable data point in what is
likely to be a wide range of human and animal behavior it
can be used as a starting point in furthering questions concern-
ing human subsistence activities and their archaeological pat-
terning. Further tests of the hypothesis advanced here should
include more actualistic and archaeological research that em-
phasizes the differential impact of large (e.g., hyaenids and ur-
sids) and small predators (i.e., canids, mustelids, etc.) on small
fauna (i.e., lagomorpha, cynomys, etc.) and their potential for
patterning within zooarchaeological collections (cf. Andrews,
1990).
Admittedly, the most difficult component of finding re-
mains influenced by human mastication is the problem of rul-
ing out other taphonomic and carnivore mastication activities.
While it may be appropriate in many instances to attribute
mastication damage to carnivores and then differentiate those
bones from other cultural activities, it should not be assumed
a priori. It is entirely possible that human mastication can be
archaeologically and paleoanthropologically identified in col-
lections of small mammal remains based on observed tooth
mark damage frequencies with minimal amounts of bone con-
sumption; especially when found in association with other cul-
turally patterned subsistence remains. What this research
suggests is that initial identifications of a canid influence on
small mammal remains may often be in error. If such is the
case, then any model seeking to discuss a range of human di-
etary choices would be strengthened by considering the impact
of humans in small faunal assemblages that do not otherwise
exhibit strong technological indications of a human presence.
Acknowledgements
This research was supported in part by a grant from the
L.S.B. Leakey Foundation. The author continues to be in-
debted to Dr K. Lupo for providing the opportunity to partic-
ipate with her research amongst the Bofi. Thanks are owed to
Alain Peneloin and Georges N’Gasse who gave project mem-
bers permission to conduct research in the Ngotto Forest Re-
serve for this collection. My deepest gratitude goes to Dr C.
Davitt, Dr V. Lynch-Holm and the late Dr V. Franceschi, at
the Electron Microscopy Center, Department of Biological
Sciences at Washington State University, for supporting the
microscopic work contained herein. Special thanks goes to
three anonymous reviewers as well as Dr M. Mondini, Dr R.
Quinlan and J. Fancher for their advice and support in improv-
ing this work, though any errors in the final product are cer-
tainly the responsibility of the author alone.
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