Tooth marks and human consumption: ethnoarchaeological mastication research among foragers of the Central African Republic

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 opportunity 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 suggests 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 paleoanthropologists 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 relation 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: [email protected] 0305-4403/$ - see front matter Ó 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.jas.2006.12.001

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 expected 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 archaeologically visible at ephemeral occupation sites. Further,

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the vagaries of invasive rodent remains that do not show evidence 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), Nunamiut (Binford, 1978, 1881), and Sioux (White, 1953, 1955) as well as two explicit exploratory studies that utilized collections of sheep in Argentina (Elkin and Mondini, 2001) and numerous 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 omnivores. However surficial the evidence may initially be for archaeological application, it suggests a means of distinguishing human and non-human accumulated small mammal assemblages. As Binford (1981:148) noted, ‘‘it is highly unlikely that a normal hominid pattern of consumption would include gnawing 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 ethnoarchaeological 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 archaeological 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 examples 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’Bae´re´ 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 semideciduous 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 season 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.), bushmeat, 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, crustaceans, 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 porcupine (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 occasionally 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 individual 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 previous 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 porcupines, 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 roasting of the porcupine, rat and mice carcasses may dry and strengthen muscle attachments, thus increasing the relative

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likelihood of teeth imprinting on a bone’s surface. Hence, differential 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 follow), 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 remote 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, researchers utilized locally available pot scrubbers as mechanical 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 relatively 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 postcollection 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 evidence of field cleaning while examining them for evidence of mastication damage. Thus, tooth marks that are visibly associated 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 shaping of the maxillary incisors when they enter adulthood.

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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 foragers 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 foragers and that not all shaped incisors are ‘fully’ shaped. In a survey of 54 adolescent and older (216 potential upper incisors) 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). Anecdotal observations indicate that when humans chew on bone (whether to gnaw on the bone itself, or in the process of removing 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 masticatory muscle force (Kieser, 1999). However, it is possible that the Bofi forager practice of tooth shaping may impact the faunal 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] impossible to define or replace .’’ (Schlosser, 2002). 3. Faunal assemblage and methods Taken as an aggregated assemblage, the Bofi faunal collection exhibits an array of damage types from field butchering, 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 describe a previously under-represented type of human processing damage. Thus, the differential distribution of multiple damage types (i.e. cutmarks, burning, etc.) across the assemblage, and their localized relationship to one another is described 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 characteristic definitions used herein are made explicit in Table 1 with comments on typical skeletal element associations. Using

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Table 1 Definitions and characteristics used in identifying tooth marked bones Type(s) of damage Description(s) Crenulated edges

Fractured edges

Tooth pits

Tooth punctures

Notches

Scoring

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) 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 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) 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) 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) 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, hindlimb 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 microscope (SEM). Both the IAW and SEM are housed in the Electron 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 consisted 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 mammals need not undergo the additional trials and tribulations of large specimen SEM work (Gilbert and Richards, 2000). Measurements 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 following Capaldo and Blumenschine (1994). 4. Damage descriptions The most obvious types of macroscopic damage on the bones resulting from human mastication activities are crenulated 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 crushing 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 surface. These elongate impressions tend to have u-shaped crosssections 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

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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 Head and necka Post-cervical axiala Forelimba Hindlimba Total tooth marked Total non-tooth marked Total assemblage a

Blue duiker

Pouched rat

Brush-tailed porcupine

Murid mice

Totals

NISP

%

NISP

%

NISP

%

NISP

%

NISP

%

26 155 33 14 228 921 1149

10.24 61.02 12.99 5.51 19.84 80.16

5 70 35 18 128 571 699

3.91 54.68 27.34 14.06 18.31 81.68

0 46 20 7 73 217 290

0.00 63.01 27.40 9.59 25.17 74.83

1 7 2 13 23 353 376

3.70 25.93 7.41 48.15 6.12 93.88

32 278 90 52 452 2062 2514

7.08 61.50 19.91 11.50 17.98 82.02

Percentages of the element groups are proportions of the tooth marked assemblage only.

layers at the point of impact. Tooth notches are often associated 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 perpetuated by a force that is driven perpendicular to the surface of the bone. As such, differences in their characteristic definitions 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 located 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 matrices (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 prerequisite 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., different enamel and bone matrices) than they are the individual actors per se. This point is made clear by Njau and Blumenschine (2006:149) with their recognition of bisected tooth marks and a specific reptilian tooth structure. The marks recorded here for the Bofi foragers, with their omnivorous tooth structure, do not differ markedly from other mammalian carnivores in the strict morphological sense of impressions left on bones with teeth. While morphological differences are unlikely to exist between predators with similar tooth structures on prey of similar sizes, it is not unreasonable to expect that the distribution, frequency, 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 qualitative 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

Fig. 1. SEM image of crushing along a broken margin of the ventral process of a pouched rat vertebrate at a magnification of 20.

Qualitatively, the blue duiker bones with human mastication 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 damage 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 diaphysis portion. On irregular or flat duiker bones, tooth marking 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

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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

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

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 increased focus on axial bones among the blue duiker remains (especially the cervical vertebrate) is likely a product of sharing and redistribution. Blue duiker are often captured in communal 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 receive 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 distributional 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 techniques 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 consumes the meat. The Bofi tend to choose the hindlimb portion 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 consumption, 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 comparatively 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, especially 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.

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Table 3 Mastication damage types within the Bofi forager faunal SEM sample SEM sample

Crenulated edgesa

Fractured edgesa

Pitsb

Puncturesb

Notchesb

Scoringb

Bones with evident field cleaning

Blue duiker (n ¼ 55) Giant pouched rat (n ¼ 26) Brush-tailed porcupine (n ¼ 21) Murid rats and mice (n ¼ 9) Total counts (n ¼ 111)

88 37 32 5 162

14 13 3 6 36

38 23 16 3 80

6 e 1 e 7

e 8 5 e 13

38 18 17 4 91

33 16 16 4 71

a b

Reported crenulated and fractured edges in numbers of individual occurrences. Reported damage is recorded as presence/absence of bones with damage type.

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 variation 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 between 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 respectively). Although field cleaning altered the surfaces of bone in the Bofi assemblage, it does not appear to have substantially 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 remains (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 duiker 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.

7. Tooth mark measurements

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Table 4 Size ranges of tooth pits measured as maximum linear dimension in millimeters Duikera Duikerb Rata Ratb Porcupinea Porcupineb Micea Miceb Total

n

Minimum

Median/mean

Maximum

Std. dev.

42 53 22 48 5 30 4 1 205

0.205 0.135 0.179 0.193 0.332 0.227 0.200 e 0.135

0.909/1.437 0.621/0.803 0.721/0.853 0.757/0.920 0.714/0.692 0.744/0.805 0.393/0.529 0.223 0.750/0.955

4.310 2.480 1.990 3.214 1.327 1.671 1.130 e 4.310

1.220 0.507 0.520 0.604 0.398 0.374 0.436 e 0.754

a Measurements of tooth pits where bone had no visible evidence of 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 the skeletal elements for small animals, this research indicates that humans and other carnivores are likely to leave similarly sized marks on small prey remains. This is not surprising as one might expect the interaction of enamel with bone to perpetuate a similar reaction once the size, strength, and density of both the bone and the actor are controlled (Binford, 1981; White, 1992). Dominguez-Rodrigo and Piqueras (2003:Table 1) reported a series of diaphyseal tooth pit measurements for a variety of carnivores where the mean length is ca. 1.5e4.0 mm and the overall weighted mean is 2.67 mm. The size range for Bofi forager tooth pits (0.14e4.31 mm) clearly coincides with the overall tooth pit range of jackals where Dominguez-Rodrigo and Piqueras (2003) report a 95% confidence interval of 0.51e4.2 mm when the diaphyseal and epiphyseal ranges are combined. The overlapping size of tooth pits between the Bofi foragers and jackals is even more remarkable as the jackal 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 than 4.0 mm in length on medium-sized bovid and equid remains (Dominguez-Rodrigo and Piqueras, 2003). The range of maximum breadth measurements for tooth scratches is provided in Table 5. Two hundred sixty-two tooth scratches were measured with the largest being less than three-quarters of a millimeter in width and the narrowest measuring less than one-tenth of a millimeter wide. Given the results of the pilot study noted earlier, many of these measurements are impacted by or derived from the field cleaning methodology. Excepting those scratches that are associated with field cleaning, the mean tooth scoring mark centers at approximately one-twentieth of a millimeter in width (n ¼ 77) while the overall range on these small prey animals does not exceed three-quarters of a millimeter. The majority of tooth scratches measured here, whether associated with field cleaning practices or not, are imperceptible without the aid of magnification. The scratch marks that do not appear associated with damage from field cleaning

Table 5 Size measurements of tooth scratches as maximum breadth in millimeters n

Minimum

Median/mean

Maximum

Std. dev.

Duiker Duikerb Rata Ratb Porcupinea Porcupineb Micea Miceb Totala Totalb

42 36 22 76 13 56 e 17 77 185

0.053 0.017 0.050 0.004 0.027 0.005 e 0.005 0.027 0.004

0.218/0.268 0.086/0.086 0.132/0.152 0.040/0.042 0.143/0.168 0.026/0.031 e 0.027/0.030 0.186/0.218 0.036/0.046

0.630 0.147 0.343 0.121 0.466 0.086 e 0.086 0.630 0.147

0.127 0.039 0.087 0.030 0.115 0.020 e 0.026 0.126 0.035

Total

262

0.004

0.057/0.097

0.630

0.108

a

a

Measurements of tooth pits where bone had no visible evidence of field cleaning. b Measurements of tooth pits where bone surface was clearly marked by field cleaning.

generally occur as single or paired marks near crenulated edges, tooth punctures and tooth pits. Previous reports have indicated a mean tooth score breadth on diaphyseal skeletal elements between 0.2 and 2.2 for a variety of carnivores (Dominguez-Rodrigo and Piqueras, 2003; Njau and Blumenschine, 2006). The average Bofi forager tooth score breadth (0.03e0.63 mm) falls at the very lower range of these means and is most similar to tooth scores left by jackals on small-sized carcasses where the 95% confidence interval is 0.19e0.73 mm (Dominguez-Rodrigo and Piqueras, 2003:Table 2). However, tooth scores left by hyaenas, baboons, bears and German Shepherds on larger prey remains all had tooth score marks that fell within the range described here for the Bofi (Dominguez-Rodrigo and Piqueras, 2003). Table 6 provides the overall size ranges of tooth punctures and tooth notch breadth and total notch ratio in this collection. The largest puncture was on the proximal end of a blue duiker tibia and the smallest puncture mark was left on a blue duiker Table 6 Size ranges of tooth punctures measured as maximum linear dimension in millimeters and tooth notch breadth dimensions n

Minimum

Median/mean

Maximum

Std. dev.

7 9 e 1 17

0.874 0.291 e e 0.291

1.018/1.949 1.360/2.191 e 2.600 1.98/2.232

4.180 4.310 e e 4.310

1.324 1.673 e e 1.444

Notch Breadth 6 Rata Ratb 4 Porcupinea 1 Porcupineb 4 Total 15 Notch ratio 15

0.714 0.482 e 0.584 0.482 0.734

1.516/1.888 1.478/1.359 3.670 1.393/2.130 1.670/1.930 1.977/2.238

4.600 2.000 e 5.150 5.150 5.255

1.447 0.654 e 2.115 1.456 1.233

Puncture Duikera Duikerb Porcupinea Porcupineb Total

a Measurements of tooth pits where bone had no visible evidence of field cleaning. b Measurements of tooth pits where bone surface was clearly marked by 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 variation 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 resembles that reported for jackal gnawed remains where the epiphyseal 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 brushtailed porcupine fibula. Although tooth notches display a large size range in breadth, the ratio of notch breadth to depth defines 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 faunal 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 porcupine (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 animal elements, where the shorter but relatively broader rat and porcupine bones are more apt to retain larger tooth marks associated with full breaching of bone walls, than are the lithe but longer blue duiker elements. Similarly, the relatively fragile 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 included 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 diaphyses may be useful in distinguishing coarse size classes of consumers though it is unlikely to indicate a specific

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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; DominguezRodrigo 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 axial 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 mastication 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; FernandezJalvo 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 assemblages 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 versus highly fragmented), the frequency of mastication across the collection (e.g., 6% of gnawed mice remains, 25% of

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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 specific 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 focus on axial elements and long bone epiphyseal portions) in association with other predators that are not etched by digestive acids and highly fragmented. Thus, while Bofi consumption 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 signature in small mammal faunal assemblages. While not suggesting 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 identical 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 considerations 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 collection, consumption of small animal remains by carnivores and hominids is likely to produce the same archaeological signature (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, ursids, felids, canids, mustelids, and other carnivores do not generally 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 concerning human subsistence activities and their archaeological patterning. Further tests of the hypothesis advanced here should include more actualistic and archaeological research that emphasizes the differential impact of large (e.g., hyaenids and ursids) 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 remains influenced by human mastication is the problem of ruling 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 collections of small mammal remains based on observed tooth mark damage frequencies with minimal amounts of bone consumption; especially when found in association with other culturally 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 dietary 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 indebted to Dr K. Lupo for providing the opportunity to participate with her research amongst the Bofi. Thanks are owed to Alain Peneloin and Georges N’Gasse who gave project members permission to conduct research in the Ngotto Forest Reserve 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 improving this work, though any errors in the final product are certainly the responsibility of the author alone. References Andrews, P., 1990. Owls, Caves and Fossils. University of Chicago Press, Chicago. Andrews, P., Nesbit Evans, E.M., 1983. Small mammal bone accumulations produced by mammalian carnivores. Paleobiology 9 (3), 289e307.

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