The role of carnivores and their relationship to hominin settlements in the TD6-2 level from Gran...

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Quaternary Science Reviews 93 (2014) 47e66

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The role of carnivores and their relationship to hominin settlements inthe TD6-2 level from Gran Dolina (Sierra de Atapuerca, Spain)

Palmira Saladié a,b,c,*, Antonio Rodríguez-Hidalgo a,b,d, Rosa Huguet a,b, Isabel Cáceres b,a,Carlos Díez e, Josep Vallverdú a,b, Antoni Canals b,a,d, María Soto a,b, Boris Santander b,a,c,José María Bermúdez de Castro f, Juan Luis Arsuaga g,h, Eudald Carbonell a,b, i

a IPHES, Institut Català de Paleoecologia Humana i Evolució Social, Unit associated with the CSIC, C/Escorxador s/n, 43003 Tarragona, SpainbÁrea de Prehistria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya 35, 43002 Tarragona, SpaincGQP-CG, Grupo Quaternário e Pré-História do Centro de Geociências (uI and D 73 e FCT), Portugald Equipo Primeros Pobladores de Extremadura, Casa de la Cultura Rodríguez Moñino. Avda. Cervantes s/n, 10003 Cáceres, Spaine Laboratorio de Prehistoria, IþDþi (Laboratory of Pre-History Research and Development), Universidad de Burgos, Plaza Misael Bañuelos s/n,09001 Burgos, SpainfCentro Nacional de Investigación sobre Evolución Humana (CENIEH), Paseo Sierra de Atapuerca, s/n, 09002 Burgos, SpaingDepartamento de Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, SpainhCentro de Investigación (UCM-ISCIII) de Evolución y Comportamiento Humanos, C/Sinesio Delgado, 4 (Pabellón 14), 28029 Madrid, Spaini Institute of Vertebrate Paleontology and Paleoanthropology of Beijing (IVPP), China

a r t i c l e i n f o

Article history:Received 13 February 2014Received in revised form28 March 2014Accepted 1 April 2014Available online

Keywords:Carnivore tooth marksHuman tooth marksCarnivore ravagingCompetitionCannibalism

* Corresponding author. Institut Català de Paleoecocial (IPHES), C/Marcel.lí Domingo s/n e Campus SescelTarragona, Spain. Tel.: þ34 977 94 30 03x3013.

E-mail addresses: [email protected], psaladie

http://dx.doi.org/10.1016/j.quascirev.2014.04.0010277-3791/� 2014 Elsevier Ltd. All rights reserved.

a b s t r a c t

Pleistocene level TD6-2 of the Gran Dolina site (Sierra de Atapuerca, Spain) is the result of anthropogenicaccumulation. Hominin groups occupied the cave as a home base, where they brought in, butchered andconsumed the carcasses of ungulates and other hominins. In this paper, we reassess the role of carnivoresin the formation and/or modification of the assemblage. We employed different methods to explore thescenario in which the TD6-2 assemblage was formed: (1) identifying the actor responsible for toothmarks; (2) determining the frequency of carnivore tooth marks and their distribution; (3) identifying theco-occurrence of modifications (butchering marks and carnivore tooth marks); (4) calculating the per-centage of change and the epiphysis to shaft ratio. Carnivore tooth marks are scarce, as is the co-occurrence of hominin and carnivore modifications. However, not all tooth marks have been attrib-uted to a particular agent due to the high equifinality between human and carnivore tooth marks. Forthese reasons, the frequency of tooth marks and the co-occurrence of modifications have been of littlehelp in interpreting the role of carnivores.

Axial skeletal remains and the epiphyses of the long bones are in large part missing. The percentage ofchange and the epiphysis to shaft ratio suggest moderate carnivore ravaging activity. Our data indicatethat the role of carnivores in TD6-2 seems to have had an impact on the original assemblage afterhominins had extracted a large amount of nutrients from the carcasses. Cannibalized hominin remainsshowed no carnivore tooth marks and have a greater presence of low survival bones compared to un-gulate remains. These findings point to a different taphonomic history suggesting that TD6-2 representsa succession of settlements having different characteristics.

� 2014 Elsevier Ltd. All rights reserved.

1. Introduction

The extent of the relationship between hominins and carnivoresin Plio-Pleistocene assemblages has been extensively explored in

logia Humana i Evolució So-ades URV (Edifici W3), 43007

@iphes.cat (P. Saladié).

the taphonomic literature. Numerous actualistic (naturalistic andexperimental) observations have established various scenariosinvolving both hominins and other large predators (e.g. Selvaggio,1994a, 1998; Blumenschine and Selvaggio, 1988; Capaldo, 1995,1997, 1998; Domínguez-Rodrigo, 1997, 1999; Egeland et al., 2004;Domínguez-Rodrigo and Barba, 2006; Espigares et al., 2013;Egeland, 2014). The results of this archaeological and actualisticresearch suggest that archaeological sites may be the product ofdifferent models of access to carcasses for hominins (hominin to

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P. Saladié et al. / Quaternary Science Reviews 93 (2014) 47e6648

carnivore, carnivore to hominin, and carnivore to hominin tocarnivore) and different levels of intra- and inter-specific compe-tition (e.g. Blumenschine and Marean, 1993; Domínguez-Rodrigoand Organista, 2007; Faith et al., 2007; Egeland, 2008). Thesemodels have been used to interpret archaeological fauna from theEarly Pleistocene, and they have made it possible to establish thesequence of different actors’ access to carcasses in order to betterunderstand the subsistence strategies of early hominins. Inferringwhich of these settings occurred during the formation of anassemblage is important but also complicated in the case of EarlyPleistocene deposits. These assemblages are often palimpsests inwhich different events, occupation types, accumulators and/ormodifying agents are accrued and overlaid; and amongwhich theremay be a greater or lesser degree of interdependence (e.g. Oliver,1994; Domínguez-Rodrigo et al., 2002; Egeland et al., 2004; Faithet al., 2007).

Level TD6-2 of Gran Dolina is a key site for understanding thesubsistence strategies and behavioral skills of hominins during theEuropean Early Pleistocene. Taphonomic studies of the macro-mammal assemblage found in TD6-2 suggest that groups of hom-inins occupied the cave as a home base during the Early Pleistocene.According to Saladié et al. (2011, 2012), the TD6-2 hominin groupswere the main accumulating agents of the large mammal remains.Modifications by carnivores have also been documented in theassemblage. In this respect, Díez et al. (1999) pronounced thecarnivore ravaging in TD6-2 as irrelevant (based on a less-studiedsample than this one). They also suggested that large felids werepossibly the first to access carcasses in the 4 to 6 weight size(Table 1). Saladié et al. (2011), however, concluded that homininshad early access to carcasses of all weight sizes, and indicated thatthe anatomic profiles with a clear shortage of axial elements maybe related to carnivore involvement in modifying the set created byhominins. This hypothesis puts carnivores as playing a larger role inthe modification of the macromammal remains.

In this paper, we explore in detail the role of carnivores in EarlyPleistocene level TD6-2, and the degree of inter-relationship be-tween the actions of hominins and the actions of carnivores in anassemblage in which anthropogenic activity predominates. Ourapproach is based on two preliminary premises in keeping withOliver’s (1994) notes: (1) firstly, taphonomic agents do not leavesignatures on all of the specimens with which they interact; (2) andsecondly, we must carefully consider any inferences drawn fromthe presence and frequency of tooth marks because not all pits,punctures or scores on a bone allow us to identify the actor as a

Table 1Grouping of taxa of TD6-2 level by sizeweight categories (list of taxa from Bermúdezde Castro et al., 1997; García and Arsuaga, 1999, 2001; Van der Made, 2001; García,2003).

Size 6 (>1000 kg) Adult Stephanorinus hundmeinensisSize 4e5 (300e1000 kg) Juvenile and infantile Stephanorinus

hundmeinensisEquidae (stenionan)cf. Bison voigstedtensisEucladoceros giuliiInfantile Mammuthus sp.

Size 3 (100e300 kg) Adult and juvenile Cervus elaphusAdult Crocuta crocutaAdult and juvenile Ursus dolinensis

Size 1e2 (10e100 kg) Homo antecessorDama nestii vallonetensisInfantile Cervidae indet.Sus scrofaCanis mosbachensisVulpes praeglacialisLynx sp.Cercopithecidae indet.

carnivore. In this context, our main aim is to establish the rela-tionship between carnivores and hominins in TD6-2 by means oftaphonomic criteria. To achieve this objective we examined severalmethods for assessing the integrity of the anatomic profile of theassemblage, the possibility of assigning tooth marks to a particularactor, the distribution of tooth marks on the bones and portions ofbone, and the co-occurrence of carnivorous tooth marks andanthropogenic modifications on the same specimens.

2. The Gran Dolina site and level TD6

In 1994, the biostratigraphic test pit reached the top of levelTD6, from which an unprecedented set of lithic tools, human andfaunal remains were recovered. Several stratigraphic studies haveestablished that level TD6 of Gran Dolina dates to the late EarlyPleistocene (Parés and Pérez-González, 1995, 1999; Falguères et al.,1999; Pérez-González et al., 2001; Berger et al., 2008; Parés et al.,2013). The TD6 sedimentary unit containing palae-oanthropological remains was named the Aurora Stratum by thefield team. The Aurora Stratum was excavated during the 1994 and1995 fieldwork season. Microstratigraphic research on the northernprofile of the level TD6 test pit revealed a bed set of at least fourrhythms of deposits in lithofacies within the Aurora Stratum(Vallverdú et al., 2001; Canals et al., 2003). This phenomenonsuggests an occupational diachrony in the formation of the AuroraStratum. However, this diachrony was difficult to quantify becauseits lateral continuity is very limited (Canals et al., 2003).

New archaeological excavation work began in 2003 in level TD6and continued until 2009. This new field survey provided an op-portunity to observe the lateral continuity of the Aurora Stratum.New human remains, macromammal fossils and lithic tools wererecovered in this new sector (w12 m2) (Bermúdez de Castro et al.,2008). Stratigraphic studies in this new area (squares G14 and G15)revealed an increased thickness of the Aurora Stratum of w45 cm.The lateral termination of the sedimentary facies of the northernprofile test pit shows interdigitation, with sedimentary gravel de-posits filling scoured surfaces. In this new excavated surface, weobserved six well-differentiated layers which correlate with theAurora Stratum defined in 1995. The Aurora stratum therefore rep-resents a condensed deposit of these layers, which could not bedistinguished in the first test pit because of the lateral variations inthe sediments (Bermúdez de Castro et al., 2008). These layers nowform part of TD6-2.

3. Methods

For this study, the data described for each specimen wereelement, taxon, size, position (right or left), age, portion and side.We established the MNE (Minimum Number of Elements), the MNI(Minimum Number of Individuals) and the %MAU (Minimum Ani-mal Units). To calculate the correlation between %MAU andmineraldensity of bones we used sheep for 1e2 weight size animals(Lyman, 1994), caribou for 3 sized animals, blue wildebeest for 4-5sized animals (Lam and Pearson, 2005), and the Suby (2006) datafor Homo antecessor. The bone surfaces were macroscopically andmicroscopically examined (OPTHEC 120). Cut marks were identi-fied on the basis of the criteria of Binford (1981), Potts and Shipman(1981), Shipman (1981), Shipman and Rose (1983) and Domínguez-Rodrigo et al. (2009). We recorded the presence/absence and lo-cations of percussion pits and adhered flakes on the surface of eachof the remains (Blumenschine and Selvaggio, 1988).

Traditionally, tooth marks have been attributed to carnivores orrodents. However, the taphonomic literature is replete withwarnings that these modifications may have been caused byhominins (Twain, 1871; Binford, 1981; Brain, 1981; White, 1992;

P. Saladié et al. / Quaternary Science Reviews 93 (2014) 47e66 49

Landt, 2007; White and Toth, 2007; Martínez, 2009; to mentionjust a few relevant examples). The interest in establishing criteriathat allow us to differentiate hominin-induced tooth marks fromthose caused by carnivores is currently giving rise to a growingbody of research (Landt, 2007; Fernández-Jalvo and Andrews, 2011;Pickering et al., 2013; Saladié et al., 2013a). Despite the methodo-logical framework now being developed, we are aware that withthe information currently available, it is still impossible tocompletely isolate the product of one agent from that of another(hominins versus carnivores). Nevertheless, during the analysis ofTD6-2 we were able to create two sets of bones modified by one orthe other of these actors from both morphological and metricalcriteria and their distribution on the bones.

Hominin tooth marks were identified based on the criteria ofSaladié et al. (2013a) and taking into consideration the descriptionspublished by Fernández-Jalvo and Andrews (2011) and Pickeringet al. (2013). The human tooth marks include furrowing,scooping-out, crenulated and saw-toothed edges, longitudinalcracking, crushing, peeling and tooth marks. Human tooth markswere determined based on their morphological features (presenceof flaking and micro-striations in scores, as was the morphology ofthe pits and punctures: crescent or angular), their location on thebones and the co-occurrence of the modifications on single bone(e.g. tooth marks associated with peeling). We noted the presenceor absence of the tooth marks on each specimen.

Non-human carnivore tooth marks (hereafter carnivore toothmarks) were also present in the TD6-2 assemblage. These signa-tures were identified from the intensity of the modifications tobones, the affected animal’s weight, and themorphology of the pits,punctures and scores. The morphological traits of the punctures(deep, multicuspid, with a bowl-shaped transversal section) andthe scores (deep, with the bottom and walls creating an irregularpath, often associated with pits with an oval or angularmorphology) (Bunn, 1981; Shipman, 1981; Blumenschine, 1995;Fisher, 1995; Domínguez-Rodrigo and Barba, 2006), along withthe presence of other severe and extensive modifications to bonesof size 3 and larger animals in the assemblage led us to infer thatthere had been carnivore activity. We noted the presence of licking,intensive pitting and scoring, and scooping-out (Binford, 1981).Bones affected by stomach acid during digestionwere also includedin this group.

We also noted the measurements of the pits, punctures andscores using the criteria of Domínguez-Rodrigo and Piqueras(2003) and Andrés et al. (2012) and compared them with experi-mental data from Selvaggio (1994b), Delaney-Rivera et al. (2009),Andrés et al. (2012), Saladié et al. (2013a; 2013b), and Sala andArsuaga (2013).

The location, segment, portion and side of all the anthropogenicand carnivore modifications on the bones were recorded(Blumenschine and Selvaggio, 1988; Blumenschine, 1995;Domínguez-Rodrigo, 1997, 1999). We used ArcGIS software toillustrate the distribution of cut marks and carnivore tooth markson limb bones. The GISmethod for counting theMNEwas originallydescribed by Marean et al. (2001). Abe et al. (2002) also used it toestablish a method for standardizing the frequency of cut marks bysurface area. Parkinson (2013) has recently incorporated the use ofSpatial Analyst ArcGIS Module tools. The Density tool (KernelDensity) is useful for identifyingwhere clusters of modifications arelocated on specific elements. Following Parkinson (2013), the exactposition of cut and carnivore tooth marks has been spatially plottedin a single layer file over a template of a complete bone, resulting ina composite record of the distribution of bone modifications foreach element. To illustrate the distribution of modifications, weused Marean’s “Bone Sorter” extension for ArcView (Marean et al.,2001; Abe et al., 2002).

To assess the role of carnivores and ravaging effect on the TD6-2assemblage, we evaluated the ratio of epiphysis to shaft fragmentsas well as the percentage of change in the epiphysis of the longbones (Blumenschine andMarean, 1993; Domínguez-Rodrigo et al.,2002). These indices assess the degree of survival of epiphysesversus shafts. The percentage of changewas calculated according tothe method described by Marean and Spencer (1991) andBlumenshine and Marean (1993) and modified based on themethod used by Domínguez-Rodrigo et al. (2002). The initial for-mula was: (MNE before ravaging e MNE after ravaging)/(MNEbefore ravaging)*100. This was established to assess the degree ofremoval of epiphyses by carnivore activity in experimental studies,in which investigators knew the initial composition of the sample.The percentage change shows the relative change of the MNE ofeach portion between pre-ravaging and post-ravaging. High per-centage change values suggest a high degree of attrition and lowaccuracy in estimating the original abundance of the elements(Marean and Spencer, 1991; Blumenschine and Marean, 1993).Domínguez-Rodrigo et al. (2002) proposed, using the total MNE forthe analysis of archaeological assemblages as equal to the pre-ravaged MNE. According to these researchers, the total number ofepiphyseal fragments in an archaeological assemblage may begreater than the assumed number, but working with completeepiphyses according to the MNE can yield a minimum estimate ofthe percentage of change.

To assess the level of ravaging we applied the ratios proposed byDomínguez Rodrigo and Organista (2007) for assemblages inwhichearly anthropic activity has been demonstrated: the ratio of(ribs þ vertebrae) to (limb bones), and the ratio of (proximalhumeri þ distal radii) to (distal humeri þ proximal radii) (from theMNE). In these ratios, the less dense elements or element portionsare the numerators and the denser elements or element portionsare the denominators, so lower values would indicate moreintensive ravaging. A ratio value of 1 would indicate that thenumber of bones present is equal to the skeletal parts in a completeskeleton, while more intense ravaging would yield lower ratios. Avariation of these proxies is proposed by Egeland (2008) to assessthe level of competition inwhich an assemblagewas formed. In thiscase, the variables used are the ratio of (axial bones to limb bones)and the ratio of (long bone epiphyseal fragments) to (long boneshaft fragments). Thus, sites in low competition areas will showhigh axial to limb and high epiphysis to shaft ratios and vice versa.

We took an archaeostratigraphic approach to the faunal andhominin remains in level TD6-2. We used the Arché Plotter soft-ware developed at the IPHES by one of authors of this study (A.C.).The three-dimensional coordinates of the faunal and human re-mains were plotted in two-dimensional projection planes. TheArché plotter enabled us to perform oblique vertical projections inrelation to the excavation plane.

Finally, we considered the actualistic (Capaldo, 1995) andarchaeological data samples from African Early Pleistocene sites(Blumenschine, 1995; Monahan, 1996; Capaldo, 1997; Domínguez-Rodrigo et al., 2002; Egeland et al., 2004) to interpret the results ofTD6-2 with regard to the co-occurrence of hominin and carnivoremodifications to limb bone specimens.

4. Results

Over the course of this study, we analyzed 4412 remains. Theassemblage is comprised of a wide variety of taxa corresponding to16 species (Table 2), including the remains of small, medium-sizedand large carnivores. However, the remains of these animals arescarce (NISP ¼ 56 and MNI ¼ 5) compared with the remains ofherbivores, as is typical in anthropogenic accumulations. Of theunidentified remains, size 3 animals are the largest group. Bone

Table 2Taxa determined in the TD6-2 level of Gran Dolina. The table shows the NISP andMNI (list of taxa from Bermúdez de Castro et al., 1997; García and Arsuaga, 1999,2001; Van der Made, 2001; García, 2003).

Taxa NISP MNI

TOTAL Homo antecessor 165 11Eucladoceros giulii 7 2Dama nestii vallonetensis 20 2cf. Cervus elaphus 293 4Cervidae indet. 271 2cf. Bison voigtstedtensis 119 4Equus (stenonian) 59 3Stephanorhinus hundmeinensis 45 2Sus scrofa 1 1Cercopithecidae indet. 2 1Mammuthus sp. 1 1Total herbivores 818 22Canis mosbachensis 17 1Vulpes praeglacialis 7 1Canidae indet. 1 e

Ursus dolinensis 9 1Crocuta crocuta 3 1Lynx sp. 4 1Carnivora indet. 15 e

Total carnivores 56 5Size 6 15 e

Sizes 4e5 503 e

Size 3 888 e

Size 1e2 560 e

Indeterminates 1407 e

Total indeterminates 3373 e


surfaces were well preserved, so the results of postdepositionalprocesses, although present, are scarce (Saladié et al., 2011), withthe exception of the black stains from manganese oxide depositsfound on 33.7% of the remains, and the cemented sedimentattached to specimen surfaces found on 28.7% of the remains.

The assemblage was comprised primarily of shaft fragmentsfrom limb bones. Low-survival elements were also present in theassemblage, but were scarce (Table 3). The correlation between the

Table 3NISP, MNE and %MAU grouped by size weight category and Homo antecessor remains.

NISP/NME/%MAU Size 6 Size 4e5 Size 3

Antler/Corn e/e/ 4/�/� 55/�/�Skull 2/2/100 23/6/100 62/5/1Mandible 6/2/50 9/6/7.6 13/7/7Isolated Teeth 13/�/� 71/�/� 52/�/�Hyoid �/�/� 1/1/1.7 �/�/�Vertebra 2/2/3 17/7/3.5 49/7/4Clavicle �/�/� �/�/� �/�/�Ribs 4/1/0.04 72/11/7.6 89/8/6Coxae 1/1/0.5 2/1/8.3 7/1/10Scapula �/�/ 4/2/16.7 6/2/20Humerus 1/1/0.5 12/6/50 34/7/7Radius �/�/ 19/4/33.3 26/7/7Ulna 2/1/0.5 6/1/8.3 8/3/30Carpal 1/1/0.08 9/�/12.5 1/�/1.7Femur 1/1/0.5 15/4/33.3 30/5/5Patella �/�/� �/�/� �/�/�Tibia �/�/� 31/11/91.7 26/7/7Fibula �/�/� �/�/� 2/2/20Talus �/�/� 2/2/16.7 2/2/40Calcaneus �/�/� 2/2/16.7 �/�/�Tarsal 1/1/0.1 1/1/1.7 4/4/8Metapodial �/�/� 44/12/50 42/12/Phalange 1/1/0.04 15/10/6.9 13/13/Long bone 1/�/� 181/�/� 461/�/Flat bone 4/�/� 118/�/� 198/�/Articular bone 2/�/� 14/�/� 10/�/�Indeterminates 1/�/� 58/�/� 3/�/�

mineral density of the bones and the %MAU (large-sized, rs.¼ 0.318,p ¼ 0.002; medium-sized, rs ¼ �0.188, p ¼ 0.237; small-sized,rs ¼ 0.142, p ¼ 0.230; H. antecessor, rs ¼ 0.365, p ¼ 0.148) shows apositive and statistically significant relationship, albeit slight, forthe large-sized weight size. The results for the medium and smallweight sizes and H. antecessor are not significant.

4.1. Cut marks and anthropogenic breakage

Saladié and colleagues in previous studies (2011, 2012) haveextensively described the anthropogenic modifications made dur-ing the butchering process. However, for the purposes of this paper,we will summarize the presence of these modifications in theassemblage in order to better understand the activity of carnivoresin the assemblage and their relationship to hominin settlements.Butcheringmodifications included cut marks and bone breakage. In13.2% of the remains some type of butchering mark isvisible � either cut marks, bone breakage, or both. These modifi-cations affected many of the taxa (including carnivores) docu-mented on the site and all weight categories. The taxonomic groupscontaining the largest number of specimens with anthropogenicmodifications were deer (n ¼ 106) and H. antecessor (n ¼ 80).

Ten percent of the recovered remains exhibit cut marks. The cutmarks have been related to various butchering activities includingskinning, defleshing, disarticulation, dismembering, evisceration,periosteum removal, and possibly tendon removal. The proximal(femora and humeri) and intermediate (radii/ulnae and tibiae)appendicular elements have a higher percentage of cut marks(38.8% and 35.1% respectively), while the distal elements have alower percentage (17.2%) (Table 4). The remains of the post-cranialaxial skeleton also often bear cut marks. 29.3% of the ribs and 9.4%of the vertebrae showed modifications produced during thebutchering process.

Anthropogenic bone breakage (4.9%) was documented on thelong and flat bones (Table 5). The bone surface damage related toanthropogenic breakage consists of percussion pits and peeling

Size 1e2 Homo antecessor Indet.

2/�/� e/�/� 16000 62/5/83.3 25/5/100 120 10/2/16.7 5/5/50 1

15/�/� 24/�/� 311/1/8.3 �/�/� e

.2 63/11/5.6 19/14/8.5 9�/�/� 3/3/11.7 1

.7 129/18/12.5 31/14/10 165/18/41.7 2/1/30 19/6/50 3/3/30 1

0 9/2/16.7 3/3/20 e

0 14/7/58.3 2/2/20 e

4/4/33.3 2/2/8.3 e

6/�/8.3 5/5/20 e

0 25/12/100 4/2/20 e

2/2/16.7 2/2/20 e

0 10/6/50 2/2/20 e

�/�/� 2/2/20 e

2/2/16.7 e e

�/�/� 1/1/10 e

3/3/8 �/�/� e

60 27/10/60 6/5/5 310.8 19/17/1.8 24/18/6.7 e

� 186/�/ �/�/� 9� 95/�/� �/�/� 9

10/�/� �/�/� 35/�/� �/�/� 1312

Fig. 1. Classic peeling on a Homo antecessor rib (a) and on a size 3 animal vertebra (band c).

Table 4NISP and frequency of elements from TD6-2 level with cutmarks. (Proximallimbs ¼ humeri and femurs; intermediate limbs ¼ radii/ulnae, tibiae and homininfibulae; and distal limbs ¼ metapodials).

Size 6 Size4e5

Size 3 Size1e2

Homoantecessor

Total

ProximalLimb

1/250

13/2748.1

22/6434.4

11/3432.5

5/771.4

52/13438.8

IntermediateLimb

0/00

18/5036

16/5429.6

8/2433.3

5/683.3

47/13435.1

Distal Limb 0/00

10/4323.2

5/4012.5

2/277.4

3/650

20/11617.2

Skull 0/20

1/205

3/585.2

3/624.8

6/2128.6

13/1637.9

Mandible 2/633.3

1/812.5

2/1118.2

1/1010

2/540

8/4020

Vertebra 2/2100

2/1711.8

4/498.2

1/631.6

5/1926.3

14/1509.3

Rib 2/450

23/7231.9

17/8919.1

12/2941.4

12/3138.7

66/22529.3

Coxa 0/10

0/20

1/714.3

1/520

1/250

3/1717.6

Scapula 0/00

2/450

2/633.3

0/90

1/33.3

5/2222.7

Phalange 0/00

3/1520

2/1315.4

3/1915.8

4/2317.4

12/7017.1


(Johnson, 1985; Blumenschine and Selvaggio, 1988; White, 1992;Pickering et al., 2013). In addition to percussion pits, the bonesalso occasionally presented conchoidal and adhered flakes. Wefound percussion breakage on 112 specimens, mainly on longbones, although it was also present in mandibles and crania of H.antecessor. Eight axial elements (Table 5) and two phalangescompleted this group. Percussion on axial items may have beencarried out for the purpose of disarticulating these elements.

Peeling was documented in 108 flat bone specimens, mainly onsmall animal (size 1e2) and H. antecessor bones. According to therecent observations of Pickering et al. (2013), the type of peelingobserved in TD6-2 is classic or general (Figs.1 and 2 a, b). Mandiblesshowed peeling on the coronoid process because of mandible andcranium disarticulation. Meanwhile, on the vertebrae and ribs itseems that this modification occurred in two different ways. Theribs showed evidence of peeling at various heights along thediaphysis. The fractures located at the neck may have occurredduring disarticulation, as well as some of the fractures on thearticular apophyses of the thoracic vertebrae. However, the peeling

Table 5NISP and percentage of bone breakage induced by hominins and carnivores.

NISP Homininsbonebreakage

% HomininsBonebreakage

Carnivoresbonebreakage

%Carnivoresbonebreakage

Long bonesHumerus 75 8 10.7 3 4Radius 69 7 10.1 5 7.2Ulna 22 5 22.7 0 0Femur 122 7 5.7 2 1.6Tibia 72 15 20.8 1 1.4Metapodial 122 11 9.0 3 2.5Total 482 53 11.0 14 2.9Flat bonesCranium 186 9 4.8 1 0.5Mandible 41 1 2.4 2 4.9Vertebra 159 17 10.7 2 1.3Rib 341 59 17.3 0 0Scapula 23 3 13 0 0Coxa 61 1 1.6 0 0Total 529 90 17 5 0.9Phalange 72 4 5.5 2 2.8

located on rib shaft fragments and on the transverse apophyses ofvertebrae was associated with the mastication and manual/oralfracture of these bones (see section 4.2.1). Such fractures on themetapodials and phalanges of H. antecessor were associated, insome cases, with small tooth marks (Fig. 2c), suggesting that theyalso occurred during consumption.

4.2. Tooth marks

There were 501 remains with tooth marks in level TD6-2. Ofthese, 159 (3.6%) specimens were attributed to anthropogenicconsumption and 224 (4.8%) to consumption by other carnivores.The remaining bones (118 specimens, 2.6%) showed tooth marksthat were not attributed to any specific agent due to the absence ofdiagnostic characteristics. This set, unfortunately, did not help us tointerpret the behavioral patterns of the actors involved in the for-mation of the assemblage and are therefore not taken intoconsideration in the following descriptions.

4.2.1. Human tooth marksHominin tooth marks were identified mainly on the axial skel-

eton specimens (vertebrae and ribs) (Fig. 2). The 1e2 and 3 weightsizes were most affected, along with H. antecessor specimens(Table 6). The types of tooth marks documented, in order of sig-nificance, were pits, punctures and scores (present in 80.6% of thebones in this group), peeling (classic or general) (29.3%), crushing(27.5%), longitudinal cracks (17.5%), furrowing (6.2%) and scoopingout (10%), and crenulated (3.7%) and saw toothed (3.7%) edges.These modifications were generally found in association with oneanother.

Peeling found in association with shallow pits and scores,crushing, furrowing, and longitudinal cracks raises less doubt aboutthe anthropogenic origin of the tooth marks (Fig. 1b, and 2aec).Fractures occur at different heights along the ribs (Figs. 1a and 2a,b), and the vertebrae display peeling on the apophyses (Fig. 1b).

Flaking on the shoulders and bottom of scores has been docu-mented (Fig. 2d). Score lengths ranged between 0.10 mm and2.73mm. Crescent-shaped pits were also found (Fig. 2e) and rangedin size between 0.43 and 6.04 mm in length and 0.15 and 4.48 mminwidth. These toothmarks were within the range (95% confidence

Fig. 2. Human tooth marks from level TD6-2. a) General peeling on a Homo antecessor rib. The rib shows scooping-out in the head and crushing on the distal end. b) Classic peelingon an H. antecessor rib. A small pit is present on the distal end associated with peeling and longitudinal cracks. c) Proximal phalange of H. antecessor. These remain shows moderatefurrowing moderate on the epiphysis and classic peeling associated with one small pit. d) Human tooth scores that show flaking at the shoulders and at the bottom. Left imageshows an archaeological example from TD6-2. Right image is an experimental tooth mark (Saladié et al., 2013a). d) Crescent-shaped human tooth pit on a size 1e2 animal.


Table 6Specimens with human tooth marks compared to NISP.

Size4e5

Size 3 Size1e2

Homoantecessor

Indeterminate

Antler 0/40

0/550

0/20

0/00

0/1600

Skull 0/250

0/620

1/621.6

0/250

0/120

Mandible 0/150

0/130

0/100

0/5 0 0/10

Hyoid 0/10

0/00

0/10

0/00

0/00

Clavicle 0/00

0/00

0/00

0/30

0/00

Vertebra 0/190

7/4914.3

9/6314.3

0/190

0/90

Rib 8/7610.5

10/8911.2

21/12916.3

5/3116.1

0/160

Scapula 0/40

2/633.3

2/922.2

1/333.3

0/10

Coxa 0/30

0/70

0/50

1/250

0/10

Humerus 0/130

0/340

0/90

0/30

0/00

Radius 0/190

1/263.8

0/140

2/2100

0/00

Ulna 0/80

3/837.5

1/425

0/20

0/00

Femur 0/160

1/303.3

2/258

0/40

0/00

Tibia 0/310

0/260

1/1010

1/1100

0/00

Fibula 0/00

0/20

0/00

1/250

0/00

Patella 0/00

0/00

0/20

0/20

0/00

Talus 0/20

0/20

0/20

0/00

0/00

Calcaneum 0/20

0/00

0/00

0/10

0/00

Carpal/Tarsal 0/270

0/140

1/175.9

0/50

0/00

Metapodial 0/440

1/422.4

0/270

3/650

0/00

Phalange 0/160

0/130

0/190

10/2441.6

0/00

Long Bone 1/1820.5

6/4611.3

5/1862.7

0/00

0/90

Flat bone 5/1224.1

15/1987.6

7/957.4

0/00

0/90

Articular bone 0/10

0/10

0/20

0/00

0/30

Indeterminate 0/00

0/00

0/00

0/00

16/13121.2


interval) of experimental human tooth marks (Delaney-Riveraet al., 2009; Andrés et al., 2012; Saladié et al., 2013a) (Figs. 3 and 4).

4.2.2. Carnivore tooth marksThe types of bone surface modifications caused by carnivores

were pits, punctures and scores (30.1%), corrosion by gastric acid(26.8%), furrowing (26.8%), crenulated edges (9.4%), scooping-out(8.9%) and/or pitting (5.6%) (Fig. 5).

Wemainly found this type of damage on bones that could not beidentified anatomically (n ¼ 115). The most abundant identifiedremains included fragments of antlers (n¼ 16) (these were affectedmainly by gastric acids) followed by ribs (n ¼ 15) and metapodials(n ¼ 10), although damage was found on bones from the entireskeleton. According to the NISP of each weight category, items fromsize 3 animals, contributed the largest number of remains withtooth marks (n ¼ 64). According to the relative frequency of eachweight category, the most affected category was size 6, and theleast affected was size 1e2 (Size 6: 9.3%; size 4e5 ¼ 7.8%; size

3 ¼ 5.3% and size 1e2 ¼ 5% with carnivore-induced modifications)(Fisher’s exact p > 0.05). No carnivore tooth marks were docu-mented on H. antecessor remains.

The tooth marks were located most frequently on epiphysesfragments, with 20.5% bearing tooth marks versus the 6% docu-mented on shaft fragments. The Fisher’s exact test (p < 0.05) thesefrequency differences are statistically significant. Of the diaphysealfragments, nine specimens were from near the epiphyses, and therest were mid-shaft (Table 7) sections. Of the flat bones, we found agreater number of coxae with carnivore-induced modificationsthan any other type of bone (Table 8).

Carnivore bone breakage was scarce in the assemblage, (n ¼ 34)and mainly affected long bone fragments (Table 5), most of whichmeasured over 30mm in length. Given their average dimensions, thepits, punctures and perforation marks seem to have been made by alarge or medium-sized carnivore. Compared to actualistic samples(Selvaggio,1994b;Domínguez-Rodrigo andPiqueras, 2003;Delaney-Rivera et al., 2009; Andrés et al., 2012; Sala and Arsuaga, 2013; Sal-adié et al., 2013a, 2013b), pit length and width on dense corticalshafts show that the TD6 carnivore pit and puncture mark samplesare similar to those documented for large carnivores such as hyenas,lions, bears and wolves, with a 95% confidence interval. The sameexercise on spongy tissue shows that the measurements are similarto those produced by medium/large-sized carnivores (Figs. 3 and 4).

4.3. Co-occurrence of carnivore- and hominin-inducedmodifications

The number of remains exhibiting concurrent hominin (cut andpercussion marks) and carnivore modifications (carnivore toothmarks) has been proposed as a method for estimating the level ofinterdependence or independence between the two agents in theformation of assemblages (Egeland et al., 2004). It is important toremember that some of the tooth marks were not attributed to anyactor. Thus, this set of specimens could be expanded with theaddition of the 18 remains on which there were butchering modi-fications and tooth marks. Another 30 remains showed cut marksand/or hominin bone breakage and hominin tooth marks.

TD6-2 contains a number of remains (n¼ 24, 0.6%) with cut and/or percussion marks and carnivore tooth marks on their surfaces.Among these, 20 specimens bear both cutmarks and carnivore toothmarks (thesemarks explicitly overlap in twocases), three specimenshave cut marks, percussion pits and carnivore tooth marks, and onespecimen has cut marks, carnivore tooth marks and carnivore bonebreakage. The toothmarks overlap the cutmarks, indicating that thecarnivore activity occurred subsequent to the hominin activity.

Most of the remains on which carnivore- and hominin-inducedmodifications coincide were from animals in the 4e5 weight size(Table 9). On the long bones, concurrency was mainly found onmid-shaft fragments (Table 10).

More often, remains with human tooth marks also featured cutmarks and/or percussion marks. These characteristics were mainlydocumented on the flat bones (Table 9). Of the 159 specimens withtooth marks, 20.7% also had cut marks and/or percussion marks,and there were a further 35 remains with peeling and hominintooth marks.

Eighteen remains had tooth marks that were caused by an un-known actor (hominins or carnivores) as well as signs of butch-ering. These remains had an anatomic and size distribution similarto the set with carnivore tooth marks (Table 9).

4.4. Percentage of change

The pattern of representation of the epiphyses in relation to theshaft according to the MNE showed a clear underrepresentation of

Fig. 3. Measurements (mean and 95% confidence intervals in mm; length: top; width: bottom) for carnivore and human tooth marks on level TD6-2 on cortical bone tissue. Theresults are compared with those of actualistic studies. Samples with fewer than 30 tooth marks were excluded in accordance with the recommendations of Andrés et al., 2012(Legend: Se ¼ Selvaggio, 1994b; De ¼ Delaney-Rivera et al., 2009; A ¼ Andrés et al., 2012; S1 ¼ Saladié et al., 2013b; S2 ¼ Saladié et al., 2013a).


the former in the TD6-2 assemblage (Table 11). The percentage ofchange, in turn, indicates a low degree of survival of epiphysesversus shafts (Table 12) for all sizes. The relationship between thetwo variables (r pearson correlation) could indicate intense modi-fication by carnivores (r ¼ �0.9177; p < 0.05).

5. Spatial distribution of carnivore activity

Level TD6-2 boasts a high density of archaeological remains. Acut and fill feature measuring one meter in diameter in whichmaterials were removed by erosion was located in the north-eastern part of the excavated grid. Faunal remains and humanbones were distributed across the entire surface (Fig. 6). Remains

with carnivore activity do not present a different horizontal allo-cation compared to other remains (Fig. 7).

Examination of the longitudinal and oblique profiles shown inFigs. 7 and 8 shows some of the characteristic features of thearchaeostratigraphic levels. The vertical distribution of faunal andhominin remains is continuous, and the highest densities of re-mains are in the full deposit. Meanwhile, specimens that showcarnivore activity are dispersed vertically. However, there are areas(in the vertical distribution) with denser carnivore activity con-centrations (top and bottom of level). This may be indicating thepeak of these animals’ activity on the faunal remains of the deposit.These remains were uncovered at a distance fromwhere the largestaccumulations of H. antecessor specimens were found, and there-fore where cannibalism events were recorded (Fig. 8).

Fig. 4. Measurements (mean and 95% confidence intervals in mm; length: top; width: bottom) for carnivore and human tooth marks on level TD6-2 on cancellous bone tissue.Length (top) and width (bottom). Legend Fig. 2. *The Homo archaeological sample has a number of 19.


6. Discussion

The TD6-2 assemblage contains a significant anthropogeniccomponent (Saladié et al., 2011, 2012), since the hominins were themost important agents in the formation of the assemblage. Taph-onomic research indicates that carnivores were also involved inmodifying the assemblage. The distribution of cut marks (Fig. 9)and the fracturing of the long bones indicate that, in the majority ofcases, hominids had first access to the carcasses of ungulates andhominins. In the case of hominin remains, the only agents involvedin their modification and consumption were other humans.Carnivore modifications and fractures occur on 4.8% of the bones;this figure could be as high as 8.1% if the remains with tooth marksproduced by an indeterminate actor (carnivore or hominin) weretaken into consideration. Blumenschine (1995) argued that, in amodel inwhich carnivore access is secondary, only 5% of the central

shafts exhibit tooth marks. Capaldo (1995) documented a higherpercentage (15%) in this regard. The scarcity of carnivore toothmarks on shaft fragments and their greater presence on epiphysesand near epiphyses portions (Fig. 10) in TD6-2 seems to support amodel in which carnivores accessed the remains once they hadbeen defleshed by hominins. This degree of prevalence is lowerthan that found in actualistic research undertaken at simulatedsites where hyenas consumed remains previously fractured withhammerstone-anvil technology (Blumenschine and Marean, 1993;Blumenschine, 1995; Capaldo, 1997). The distribution of the toothmarks on the long bones shows a clear pattern that suggests thatthese animals were more active at the ends of bones. Anthropo-genic bone breakage is also more frequently found than breakagecaused by carnivores (Fig. 11) and the latter is found in 42.8% ofcases in the near epiphysis portions. According to Oliver (1994), thescarcity of bones fractured by carnivores suggests that hominins

Fig. 5. Carnivore tooth marks present on several remains from TD6-2.

Table 8Frequency (%) of non long bones with carnivore tooth marks.

Size 6 Size4e5

Size 3 Size1e2

Homoantecessor

Indeterminates Total

Antler 0 0 14.5 0 0 5 7.2Skull 0 8.3 3.1 0 0 0 2.1Mandible 0 0 9.1 30 0 0 9.8Vertebra 0 11.8 0 3 0 0 2.5Clavicle 0 0 0 0 0 0 0Ribs 25 12.5 5.6 0 0 0 4.4Coxae 0 50 14 40 0 0 22.2Scapula 0 25 0 0 0 0 4.3Carpal/Tarsal/

Sesamoide0 10.7 11.8 17 0 0 10.8

Phalange 0 13.3 7.7 11 0 0 6.9Flat bone 25 10.2 4 4 0 0 5.9Indet. 0 6.9 0 0 0 3.2 3.3Total 8 10.3 5.5 4 0 3.3 4.5

Table 9Remains with carnivore tooth marks and cut and/or percussion marks.

Size 6 Size 4e5 Size 3 Size 1e2 Homoantecessor

Indet. Total

Carnivore tooth marks D butchered marksSkull e 1 1 e e e 2Ribs e 3 1 e e e 4Scapula e 1 e e e e 1Humerus e 1 1 e e e 2Ulna 1 e e e e e 1Femur e 1 e e e e 1Tibia e 1 1 e e e 2Metapodials e 1 e e e e 1Long Bones e 2 e 2 e e 4Flat Bones e 5 1 e e e 6Total 1 16 5 2 e e 24Indeterminates Tooth marks D butchered marksRibs e e 1 1 e e 2Humerus e e 1 1 e e 2Radius e e e 1 e e 1Ulna e e e 1 e e 1Tibia e 1 e e e e 1Long bone e 1 1 1 e e 3Flat bone e 2 3 e e 5Indet. e 1 e e e 2 3Total e 5 6 5 e 2 18Hominin tooth marks D butchered marksSkull � � � 1 � e 1Scapula � � � 1 1 e 2Ribs � 4 5 10 5 e 24Vertebrae � � 3 1 6 e 10Coxa � � � � 1 e 1Radius � � 1 � 2 e 3Ulna � � � 1 � e 1Femur � � � 1 � e 1Tibia � � � � 1 e 1Fibula � � � � 1 e 1Metapodial � � � � 3 e 3Phalange � � � � 5 e 5


were the responsible for most of the accumulation, and they did notleave many whole, fresh bones due to the intensive exploitation ofcarcasses.

Furthermore, to assess the frequency and distribution of carni-vore tooth marks in TD6-2, it is important to remember that thereare also tooth marks caused by hominins, as well as a set of boneswith tooth marks of indeterminate origin. The morphology of hu-man tooth marks can sometimes be similar to those produced byother actors and may overlap with measurements of other carni-vore tooth marks (Delaney-Rivera et al., 2009; Saladié et al., 2013a).These similarities give rise to difficulties in isolating the product ofone agent or another. For 23.4% of tooth marks recorded, it was not

Table 7Frequency (%) of the distribution of carnivore tooth marks on long bone portions.The hominin bones not show carnivore tooth marks.

Size 6 Size 4e5 Size 3 Size 1e2 Total

Prox. Epiph. 0 20 25 0 21.4Prox. Epiph.þShaft 0 0 0 0 0Shaft 40 6.6 5.1 7.4 6.1Distal Epiph.þShaft 0 0 0 0 0Distal Epiph. 0 2 33.3 0 16.6Epiph. indet 0 0 50 0 14.3Whole 0 0 0 0

Long bone � � 2 2 � e 4Flat bone � 2 4 2 � e 8Indet. � � � � � 3 3

Table 10Distribution of portions of long bones containing butchering marks (cuts and/orpercussion marks) and carnivore tooth marks.

Size 6 Size 4e5 Size 3 Size 1e2 Total

Epiphyses e 1 e e 1Shaft e 5 1 2 8Near epiphyses 1 e 1 e 2Total 1 6 2 2 11

Table 11Ratio of epiphyses: diaphysis of the long bones from the TD6-2 assemblage.

NISPEpiphyses

NISPDiaphyses

Ratio Epiphyses:Diaphyses

Size 4e5 22 293 0.08Size 3 19 611 0.03Size 1e2 14 260 0.05Homo antecessor 6 17 0.35Total 61 1186 0.05

Table 12Percentage of change of long bones from the TD6-2 assemblage.

Percentage of change Size 4e5 Size 3 Size 1e2 Homo antecessor

Humerus 100 71.4 100 83.3Radius 87.5 85.7 85.7 100Femur 80 70 75.0 100Tibia 90.9 100 83.3 100Metapodial 75 79.2 90 40.0Total 85.9 81.6 83.8 75


possible to determine the actor. The fact that human tooth markshave been identified in the TD6-2 assemblage makes it necessary toexercise extreme caution in interpreting the tooth marks docu-mented on remains, in terms of both attributing carnivore ethologyand using tooth mark frequency to make inferences about thebehavior of hominins in an assemblage which contains the signa-tures of both agents. Thus, our estimate of the frequency of modi-fications produced by carnivores in TD6-2 is approximate and,without considering other features, we cannot categorically inferthe sequence of access or the level of competition between the twoagents for the same carcasses. There are two reasons for this: first ofall, we cannot assume that the carnivores are solely responsible forthe tooth marks and secondly, without solving the current prob-lems of equifinality between human and carnivore toothmarks, it isimpossible to completely isolate the signatures of the two agents. Amistaken identification of an actor’s tooth marks can result in over-or under-estimating the role that carnivores played in the forma-tion of an assemblage. We therefore, in addition to evaluating the

Fig. 6. General distribution of archaeo-paleontological remains from the excavated area of leof the following figures (Fig. 7 and 8).

frequency and distribution of carnivore toothmarks, explored othercomplementary methods for evaluating the role of these animals inthe assemblage.

There have been several researchers who have proposed othermethods for determining the relationship between hominins andcarnivores in assemblages in which the signatures of both agentsare present. These methods include: 1) the degree of co-occurrenceof hominin and carnivore modifications (Egeland et al., 2004); and2) anatomic profiles (Bunn, 1986; Marean and Spencer, 1991;Marean et al., 1992; Blumenschine and Marean, 1993; Capaldo,1995, 1998; Domínguez-Rodrigo et al., 2002; Domínguez-Rodrigoand Organista, 2007; Egeland, 2008).

The percentage of long bones in TD6-2 with cut and/or per-cussion marks as well as carnivore tooth marks is low (1.5%), and iseven lower on the flat bones (0.9%). However, there is a markedincrease in the co-occurrence of the two modifications in scapulae(4.2%). Compared to the frequency of specimens with tooth marksand the distribution of tooth marks on the remains of the twoactualistic samples published by Capaldo (1995), TD6-2 manifests amodel in which hominin and carnivore activity on the same car-casses is relatively uncommon. The same is true when the fre-quency of concurrent hominin and carnivore modifications onbones in level TD6-2 is compared to African Early Pleistocenearchaeological samples (Table 13 and Fig. 12). Level TD6-2 has afrequency of co-occurence similar to that found at the Olduvai BedII sites, ST Site Complex at Peninj, and SwartkransMember 3, wherethe hominin and carnivore signatures exhibit clear independence(Bunn and Kroll, 1986; Monahan, 1996; Domínguez-Rodrigo et al.,2002; Egeland et al., 2004). We ran a correspondence analysis toanalyze the similarities and/or differences in the co-occurence ofmodifications made by hominins and other carnivores for certainAfrican Early Pleistocene archaeological samples and the TD6-2sample. Correspondence analysis reflects the proximity of allthese sites. However, we can see that TD6 is more closely related toSwartkrans Member 3 and HWK East levels, which are the as-semblages that show a higher level of independence betweencarnivore and hominin activity. If we observe the density distri-bution of cut marks and carnivore tooth marks in different portionsof the long bones, there is limited overlay in specific areas. Fig. 13

vel TD6-2. Lines AA ‘and BB’ indicate the area that have been made vertical projections

Fig. 7. General profile (EeW) of level TD6-2.

Fig. 8. Oblique profile of level TD6-2 showing distance between greatest accumulations of H. antecessor remains and faunal remains with carnivore activity.


shows this graphically as most of the tooth marks are located closeto epiphysis portions and epiphyses, while the higher concentra-tions of cut marks are in mid-shaft portions. The low coincidence ofhominin and carnivore modifications may be due to this distribu-tion and the scarcity of epiphyses in the assemblage.

The lack of epiphyses is related to the anatomical profile of TD6-2, which shows a greater prevalence of high-survival elements. Thischaracteristic is common in faunal assemblages (Lyman, 1994) andmay have different causes, leading (again) to problems of equifin-ality (Marean and Spencer, 1991). These reasons may be related to:1) mechanical and chemical post depositional processes (e.g. fluvialtransport, bone weathering, trampling, microbial attack or chemi-cal attrition); 2) decisions made by hominins when capturing andtransporting carcasses; 3) carnivore ravaging of scraps left behindby hominins; or 4) a combination of the activities of several of theseagents or processes.

The ratio of %MAU to bone mineral density has shownthat � despite the bias present � there is no relationship betweenthese two variables. This suggests that the bias is not related topostdepositional processes. The presence of well-preserved fragileand low density bones, such as those of hominin infants, suggeststhat the assemblage has been excellently conserved. Post-

depositional modifications are scarce, which is why we cannotassume that they had a preeminent role in the attrition of theassemblage and in any modifications to the original anatomicprofile. These features lead us to discuss the role of hominins andcarnivores during the formation of the deposit. The presence of afew ribs and vertebrae of animals of all weight sizes indicates thathominins transported the axial segments to the cave on at leastsome occasions (Saladié et al., 2011). The scarcity of epiphyses oflong bones may provide some insight into this disjunctive, as thedisappearance of the ends of the long bones cannot be due to se-lective transport by hominins and is consistent with a pattern ofcarnivore post-ravaging (Blumenschine and Selvaggio, 1988;Marean et al., 1992; Blumenschine and Marean, 1993;Blumenschine, 1995; Capaldo, 1998; Marean and Cleghorn, 2003;Faith and Behrensmeyer, 2006).

The results of the percentage of change analysis of the long boneand epiphysis to shaft ratio indicate a less-close relationship e oneof independence between the actions of hominins and the actionsof carnivores in the assemblage. Simulated sites with hammer-stone breakage of carnivore bones that were mainly disturbed byspotted hyenas revealed that these carnivores focused on theconsuming the epiphyses and low-survival bone fragments

Fig. 9. Composite plot for cut marks on long limb bones of TD6-2 assemblage.


(vertebrae, ribs, scapulae, coxae, carpal/tarsal and phalanges) (Bunnand Kroll, 1986; Blumenschine and Selvaggio, 1988; Marean andSpencer, 1991; Marean et al., 1992; Blumenschine and Marean,1993; Capaldo, 1995; Marean and Cleghorn, 2003; Faith et al.,2007). After an intensive anthropogenic butchering process, theseportions retain the majority of those nutrients that are attractive toanimals such as hyenas or large canids.

The result, as is apparent from the percentage change, would bea heavy loss of low-survival elements, altering the originalanatomic profiles and therefore pointing to a high interrelationbetween the two agents that modified the same elements. In thisscenario, the carnivore activity was superimposed over homininbehavior. The scarcity of axial skeleton components might then berelated to hominins’ selective transport of meat, and certainly to

Fig. 10. Composite plot for carnivore tooth marks on long limb bones of TD6-2 assemblage.


the superimposed activity of bone-crushing carnivores such ashyenas and/or large canids. The ratios of axial bones to limb bonesand (proximal humerus þ distal radius) to (distal humerusþ proximal radius) give values of 0.28 and 0.44 respectively, whichplace TD6-2 in Stage 2 ravaging. This represents moderate ravaging

in which, for at least half the bones of each part (axial and softportions/elements) the numbers have been reduced by half(Domínguez-Rodrigo and Organista, 2007: 210). Although evidenceof carnivore activity suggests a moderate to high level of compe-tition scenario at the site (Blumenschine and Marean, 1993;

Fig. 11. Percentage of long bones with green breakage associated with percussion pits or notches from tooth marks.


Domínguez-Rodrigo and Organista, 2007; Egeland, 2008), thehominin processing of ungulate and hominin carcasses was inten-sive, suggesting that these activities were conducted in an envi-ronment characterized by little competition with other predators.Rodríguez-Gómez et al. (2013) recently modeled trophic resourceavailability at the end of the Early Pleistocene at the Sierra deAtapuerca, using the TD6-2 record. They concluded that, during theformation of TD6-2, the Sierra de Atapuerca was a rich ecosystem,with sufficient variety of prey not only to support the predators andscavengers found in the archaeological record (e.g. Canis mosba-chensis, Ursus dolinensis, Crocuta crocuta, Lynx sp. and H. antecessor),but also to ensure the survival of other large predatorse such as theEuropean jaguar Panthera gombaszoegensis and sabre-toothed catewhich have not been documented in the assemblage. Meat re-sources were abundant and competition was moderate to low inlevel TD6-2 of Atapuerca (Rodríguez-Gómez et al., 2013). However,for hominins, this probably would not preclude the risk of theft oreven injury and death in encounters with confrontational scaven-gers (such as spotted hyenas) or top-level predators (large felids).Monahan (1998) noted that the risk of encountering large carni-vores is a limiting factor for forager hominins, which significantlyalters the costs of the butchering process, transport and benefits ofthe carcasses of ungulates. In this context, Gran Dolina was used asa refuge to which hominins transported carcasses and theiranatomical parts to diminish the possibility of competition with

Table 13Summary of limb bone NISP and bone surface modification percentages for African Ea(Blumenschine, 1995; Capaldo, 1997); ST Site Complex (Domínguez-Rodrigo et al., 2002;Swartkrans Member 3 (Egeland et al., 2004) summarised by Egeland et al., 2004:350.

NISP Tooth marks and cut marks Tooth

n % n

FLK Zinj (Olduvai Bed I)Size Class 1e4

731 102 14.0 125

ST Site Complex (Peninj)Size Class 1e6

154 2 1.3 2

BK (Olduvai Bed II)Size Class 1e4

636 1 0.2 1

MNK Main (Olduvai Bed II)Size Class 1e4

313 1 0.3 0

HWK East Levels 1e2 (Olduvai Bed II)Size Class 1e4

113 2 0.6 1

Swartkrans Member 3Size Class 1e4

519 9 1.7 3

Gran Dolina TD6-2 level SizeClass 1e5

412 6 1.5 2

other predators during processing and consumption. There isgenerally a higher level of competition in open habitats than inclosed spaces such as woodland habitats, since e among othercharacteristics e in open habitats carnivores benefit from greatervisibility (Blumenschine, 1986; Domínguez-Rodrigo, 2001; Faithand Behrensmeyer, 2006; Faith et al., 2007; Egeland, 2008) of car-casses and of scavenging vultures approaching the carcasses(Palmqvist et al., 2011). A karstic context such as the Sierra deAtapuerca could therefore provide multiple refuges in which thelevel of competition with other predators would be lower and itwould even be possible to control the access of other carnivoresduring hominin occupations.

According to the records provided by Egeland (2008), in order tointerpret the level of competition, we must consider that low orhigh levels of competitionwill be different for different ecosystems,and that the time span sampled can be problematic in palimpseststhat may contain several tens or hundreds of years. At this point it iseasy to overlap (and mix) different types of occupations, which arehard to isolate. TD6-2 is the result of a sequence of events(Vallverdú et al., 2001; Canals et al., 2003) andmay therefore be theresult of a mixture of strategies, proficiencies and/or settlementtypes. Binford (1984) reported that the same sites could be occu-pied successively by Nunamiut for different purposes (as residen-tial, logistic or butchering sites, for example). These types of usageare difficult (if not impossible) to isolate in the Early Pleistocene

rly Pleistocene archaeological samples and TD6-2 sample. Data sources: FLK Zinj,Egeland et al., 2004); BK, MNK Main and HWK East Levels 1e2, (Monahan, 1996);

marks and percussion marks Tooth marks and cut and/or percusion marks

% n %

17.1 184 25.2

1.3 4 2.6

0.2 2 0.3

0.0 1 0.3

0.3 3 1.0

0.6 12 2.3

0.5 6 1.5

Fig. 12. Correspondence Analysis plot of co-occurrence of hominine and carnivore modifications for African Early Pleistocene archaeological samples and TD6-2 sample surfacemodification percentages (Data sources: ST Site Complex ¼ Domínguez-Rodrigo et al., 2002; Egeland et al., 2004; BK, MNK Main and HWK East Levels 1e2 ¼ Monahan, 1996;Swartkrans Member 3 ¼ Egeland et al., 2004; summarised by Egeland et al., 2004:350).Eigenvalue axis 1 ¼ 92.7). P þ T ¼ Specimens with least one percussion marks and least onecarnivore tooth marks; C þ T ¼ Specimens with least one cut marks and least one carnivore tooth marks; Cþ/or P þ T ¼ ¼ Specimens with least one percussion mark and/or one cutmarks and least one carnivore tooth marks).


palimpsests in which the behavior or type of occupation whichoccurred most frequently is more easily observed. However, in thisrespect, the set of cannibalized remains of H. antecessor found inTD6-2 can assist in making these interpretations.

The set of remains of the bodies of hominins are the result ofseveral events (Carbonell et al., 2010) but this set generally seems toshow a different biostratinomic history if we isolate this taxon fromthe other animals present in the assemblage. There are few carni-vore tooth marks on the bones of ungulates, but there are no suchtraces on hominin remains. The percentage change is lower on theremains of this taxon and the ratio of epiphyses to diaphyses ishigher, supporting a hypothesis of less (or no) carnivore activity onthese bodies. In the case of hominin remains, we observe that someof the more high-nutrient-content bone portions have survivedafter having been discarded by hominins. Large carnivores attackhuman bodies in a similar way to how they approach the carcassesof other animals, resulting in the disappearance of the less denseportions made up of cancellous tissue and fat (Haglund et al., 1988;Horwitz and Smith, 1988).

To measure the degree of competition, Egeland (2008) proposedto use the ratio epiphysis to shaft versus axial to limb ratio. Whenwe apply these taphonomic proxies to TD6-2, we see the creation oftwo distinct sets (Fig. 14). One group is made up of the differentweight size groups. In the other group, H. antecessor appears to beisolated. This would indicate that at least some of the ungulateremains were accumulated in an environment of moderatecompetition, but the bodies of H. antecessor (with some ungulates)were accumulated in an environment with low or no competition.The presence of very fragile and low-density elements ofH. antecessor infants, in addition to the presence of some completeor nearly complete vertebrae and fragments of ribs with anthro-pogenic fractures seems to suggest that these elements werescarcely disturbed by carnivores once they had been rejected by thehominin groups.

The archaeostratigraphic profiles of the archaeopaleontologicalremains show that the highest densities of materials with carnivoreactivity are distanced from the remains of H. antecessor. Unfortu-nately, the current number of remains and the data available to datedo not point to a solid explanation for this phenomenon, although itmay be related to the different types of occupation that occurredduring the formation of level TD6-2 and during the cannibalismevents. According to the latest research, the cases of cannibalismthat occurred in TD6-2 may be associated with times of abundantfood resources (Rodríguez-Gómez et al., 2013) and an attempt toprotect the group’s catchment territory (Saladié et al., 2012), whichsuggests that intraspecific competition may have been high. Thiswould be related to more a higher density of establishments in theterritory and therefore to longer occupations in Gran Dolina cave.Carnivores would have been present in the cave afterward theseperiods and at times when hominins were absent from the site.

7. Conclusions

Taphonomic data make clear that hominins played the majorrole in the formation of the faunal assemblage from TD6-2. Thetooth marks in the assemblage can be attributed to carnivores andhominids. It is possible to assign these tooth marks to one actorrather than another (carnivores versus hominids), although thisoften involves problems of equifinality. Thus, inferences about theinteraction between carnivores and hominins based on the pres-ence and frequency of tooth marks must be made with caution asthey may be due to a false dichotomy (all cut marks were made byhominins, but not all tooth marks can be assigned to carnivores).There is a growing need to add to the body of taphonomicknowledge through actualistic work undertaken from a uniformi-tarian perspective. These studies are useful in describing andidentifying different types of modifications. However, theseresearch frameworks also allow us to assume greater levels of

Fig. 13. GIS density analysis results of cut and carnivore tooth marks distribution performed on composite cut and tooth marks plots for limb bones from all ungulates elements(Blue tones indicate the concentrations of cut marks; brown and green tones indicate concentrations of tooth marks; Dark blue mark the areas in which the highest densities havebeen found).(For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)


equifinality between marks produced by different agents (see forexample the research of Domínguez-Rodrigo et al., 2009 in relationto the morphological characteristics of cut marks and trampling).Our study has revealed the existence of problems (sometimes

severe) in evaluating the frequency and distribution of toothmarks,and hence in assessing their co-occurence with butchering marks.

It has therefore been necessary to support the evidence from thefrequency and distribution of carnivore tooth marks with other

Fig. 14. Plot for the relationship between axial-to-limb and epiphysis-to-shaft ratios from TD6-2 sample.


taphonomic methods which are related to anatomical profiles, inorder to assess the level of competition, ravaging and assemblageformation. All the taphonomic approaches that we have exploredindicate clearly that, in the assemblage, carnivore access to remainswas secondary. We did, however, observe conflicting resultsregarding the intensity of their activity with respect to the co-occurence of modifications, anatomical profiles and the survivalof very fragile elements, and what would be expected in a ravagedassemblage. This trait is undoubtedly linked to the nature of theTD6-2 palimpsest and the overlapping of events having differentcharacteristics. Taphonomic proxies indicate that carnivores playedan important role in the destruction of the bones and bone portionsof ungulates. In contrast, the cannibalized bones of H. antecessorhave not been disturbed. No carnivore tooth marks have beenobserved on the surface of these bones, and the axial bones andepiphyses show higher survival when compared to the other ani-mals we documented. Besides, the fact that bones as fragile asvertebrae or the scapulae of infants were conserved supports theidea that carnivore activity did not occur on this taxon. To sum-marize, hominins processed and consumed the animal resourceson-site first, in a context of low competition with other predators.Meanwhile, carnivores consumed the scraps abandoned by homi-nins in a setting of low to moderate intra- and interspecificcompetition. Cannibalism at TD6-2 is related to the protection andexpansion of a catchment area. The taphonomic data presented inthis paper support the thesis that these events were associatedwith longer occupations. These two factors suggest that canni-balism occurred in a setting in which there was some level ofintraspecific competition. These traits put the European EarlyPleistocene hominins at the top of the food chain and indicate thatthey could control the animal resources evenwhen carnivores werearound. A confined space such as the karstic complex of the Sierrade Atapuerca could be a refuge that was suitable for butchering andconsuming the prey, by reducing the level of competition withother predators during the periods when the site was occupied.

Acknowledgments

We are deeply grateful to the Atapuerca research team and theparticipants in the fieldwork for that project. We thank the editorand two anonymous reviewers for their comments on the

manuscript that have greatly improved the new version. We thankDr. Charles Egeland for providing helpful comments on a previousversion of the manuscript. We are grateful to Dr. Jennifer Parkinsonso she offered to us the Marean GIS pack. We appreciate profuselythe help made by our colleague Juan Ignacio Morales and theircomments and discussion. This research was supported by theMinistry of Economy and Competitiveness (MINECO) of SpainGovernment, project n� CGL2012-38434-C03-03, and CatalonianGovernment, project n� SGR 2009-188. Funding for the fieldworkcame from Cultural and Tourism Council of Castilla y León andAtapuerca Foundation. A. Rodríguez-Hidalgo is the beneficiary of apredoctoral research fellowship (FPI) from the MINECO (CGL2009-12703-C03-02). M. Soto is the beneficiary of a predoctoral researchfellowship (FI) from AGAUR (FI-DGR 2013, Agaur). B. Santander is abeneficiary of a predoctoral fellowship from CONICYT-AdvancedHuman Capital Program (Becas Chile 2011).

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The role of carnivores and their relationship to hominin settlements in the TD6-2 level from Gran...

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