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Journal of Archaeological Science xxx (2010) 1e10

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Journal of Archaeological Science

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The hunted hunter: the capture of a lion (Panthera leo fossilis) at theGran Dolina site, Sierra de Atapuerca, Spain

Ruth Blasco a,*, Jordi Rosell a, Juan Luis Arsuaga b,c, José M. Bermúdez de Castro d, Eudald Carbonell a,e

a IPHES (Institut català de Paleoecologia Humana i Evolució Social), Unidad Asociada al CSIC, Àrea de Prehistòria, Universitat Rovira i Virgili,Plaça Imperial Tarraco, 1, 43005 Tarragona, SpainbDepartamento de Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, SpaincCentro de Investigación (UCM-ISCIII) de Evolución y Comportamiento Humanos, C/Sinesio Delgado, 4 (Pabellón 14), 28029 Madrid, SpaindCENIEH (Centro Nacional de Investigación sobre Evolución Humana), Avenida de la Paz 28, 09004 Burgos, SpaineVisiting professor, Institute of Vertebrate Paleontology and Paleoanthropology of Beijing (IVPP)

a r t i c l e i n f o

Article history:Received 22 December 2009Received in revised form15 March 2010Accepted 17 March 2010

Keywords:Subsistence strategiesHuntingCarnivore usePanthera leo fossilisGran DolinaSierra de AtapuercaMiddle Pleistocene

* Corresponding author. Tel.: þ34 977 257 882; faxE-mail address: [email protected] (R. Blasco).

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a b s t r a c t

Many Pleistocene caves and rock shelters contain evidence of carnivore and human activities. For thisreason, it is common to recover at these sites faunal remains left by both biological agents. In order toexplain the role that carnivores play at the archaeological sites it is necessary to analyse several elements,such as the taxonomical and skeletal representation, the age profiles, the ratio of NISP to MNI, theanthropogenic processing marks on the carcasses (location and purpose of cutmarks and burning and bonebreakage patterns), carnivore damage (digested bones, location and frequencies of toothmarks and bonebreakage), length of the long bones, frequencies of coprolites and vertical distribution of the faunalremains, inter alia. From this, the documentation of carnivores in a faunal assemblage with a clearanthropogenic component can be understood from three main phenomena: (1) the carnivores as accu-mulators and the use of the site as a den; (2) carnivores as scavengers of hominid refuse and; (3) carnivoresas hominids’ prey. Of these three phenomena, the last one is the least documented at the Middle Pleis-tocene sites. From this perspective, here we present the case of the anthropogenic use of a lion (Pantheraleo fossilis) from level TD10-1 of Gran Dolina (MIS 9, Sierra de Atapuerca, Burgos, Spain). The lion boneremains show signs of direct interaction between this big cat and human groups that occupied Gran Dolinain these chronologies. From this perspective, the aim of this paper is to contribute to the knowledge of therole developed by large carnivores in the anthropogenic contexts and to provide data on human use ofthese predators at the European Middle Pleistocene sites.

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1. Introduction

The archaeological excavations carried out at the Middle Pleisto-cene sites show the occupation of karstic areas by hominids andcarnivores. For this reason, the recoveryof faunal remains left by bothbiological entities is common at these sites (both the bones leftbehind from their activities and/or their own skeletal remains).Manycarnivores regularly use caves as a breedingplace or as a refuge. Thereare several studies that allow us to recognise both current andarchaeological dens (Sutcliffe, 1970; Haynes, 1980, 1983a; Hill, 1984;Blumenschine, 1985, 1986a, 1988a, 1988b; Clot, 1987; Cruz-Uribe,1991; Domínguez-Rodrigo, 1994, 2001; Selvaggio, 1994; Fosse,1996; Villa and Soressi, 2000; Villa et al., 2004; Michel, 2005; inter

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alia). However,wemust bear inmind thatnot all carnivores behave ina similar way or have the same impact on the carcasses. Each specieshas its ownethologyandphysical characteristics,which influence theaccumulations that they produce and the intensity with which theyact on bone remains. Even so, a series of general characteristics areidentified in all assemblage generated by non-human predators. Ingeneral, the age profiles of the prey tend to be attritional (with apredominance of young and senile individuals over adults). Theskeletal representation of the prey is highly variable with a domi-nance of limb bones (except for carpal and tarsal bones which can beswallowed during the consumption process) and some elements ofthe axial and cranial skeleton (Binford, 1981). The proportionbetween the cranialepostcranial skeleton of the prey decreasesaccording to the accumulating carnivore and to the size of theungulates, so the larger the prey, the lower the capacity of thecarnivore to transport the skull (Cruz-Uribe, 1991). Furthermore,coprolites, skeletal remains belonging to other carnivores, deciduous

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teeth (in the case of hyena maternity dens) and digested bones areoften retrieved at these assemblages (Cruz-Uribe, 1991; Villa et al.,2004; inter alia). Some authors, such as Fosse (1994) or Villa et al.(2004), also observed differences related to the bone fragmentationbetween the carnivore sites and anthropogenic sites. According tothese authors, the bones recovered in the assemblages generated bycarnivores usually are less fragmented and, therefore, the propor-tions of identifiable elements are much higher.

However, the carnivores are not only accumulators. The smellsfrom the remains left in the human camps are attractive for manypredators and, therefore, it is common for these carnivores to act asscavengers in search of potentially consumable elements (Binford,1981). Different observations and experimental reproductionshave been made with wild and captive animals, attempting todocument the carnivore damage on the faunal assemblages gener-ated by human groups (Sutcliffe, 1970; Klein, 1975; Bunn, 1986;Bunn et al., 1980; Bunn et al., 1988; Blumenschine, 1986a, 1986b;Marean et al., 1992). All of these studies coincide with the fact thatscavengers have a predilection for epiphyses of limb bones and for

Fig. 1. Location, composite stratigraphic profile of deposits and levels dating at the Gran Dolina

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elements of the axial skeleton. The modifications on theseanatomical portions are so intense that in many cases they makethemdisappear. On the contrary, the diaphyses, which are generallyhighly fractured during the processing and anthropic consumption,show very little impact. This duality is due to the fact that boneswith a high content of spongy tissue (epiphyses and flat bones)contain resources (fat), which are difficult to reach for the hominidswithout the appropriate technology, such as boiling (Oliver, 1993).However, during this process, carnivores do not only destroy thebones, but they can also affect the original position of the remains,by carrying out important remobilisations and significantly alteringthe spatial distribution left by human groups (Binford et al., 1988).

Nevertheless, the presence of carnivores and their damage inthe anthropogenic context, do not only respond to the scavengingactivities or to the use of the site as den. Carnivores can also bea potential prey for human groups because they can offer a series ofusable resources (skin, tendons, meat, marrow, etc.). As withthe herbivores, the processing of the carnivores is identified by thepresence of cutmarks, the intentional bone breakage and/or the

site from Parés and Pérez-González (1999), Falguères et al. (1999) and Berger et al. (2008).


Table 1NISP, MNE and MNI by ages from TD10-1 faunal sample of Gran Dolina.

TD10-1 Taxa NISP MNE MNI MNI by ages

neo. inf. juv. ad. sen.

Ursus arctos 3 2 1 1Canis lupus 10 6 2 1 1Vulpes vulpes 16 13 2 1 1Panthera leo 17 15 1 1Lynx sp. 1 1 1 1Hystrix sp. 2 1 1 1Stephanorhinus

cf. hemitoechus52 9 2 1 1

Equus ferus 260 62 9 2 3 3 1Equus hydruntinus 12 5 2 1 1Sus scrofa 1 1 1 1Cervidae 121 24 2 1 1Megaloceros giganteus 1 1 1 1Dama dama clactoniana 2 2 1 1Cervus elaphus 762 232 9 1 1 6 1Bison sp. 144 55 5 1 1 2 1Hemitragus bonali 5 5 1 1Capreolus capreolus 3 3 2 1 1Erinaceidae 11 8 1 1Leporidae 329 167 12 1 11Aves 58 40 3 3Ichthyofauna 1 1 1 1Total 1811 651 60 1 9 7 40 3

R. Blasco et al. / Journal of Archaeological Science xxx (2010) 1e10 3

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burning patterns of their remains. Although archaeological exam-ples exist in some assemblages of the first half of the Upper Pleis-tocene (Auguste, 1991,1995; Stiner, 1994; Arribas et al., 1997; David,1997; Tillet, 2002), the use of these animals is not common at theMiddle Pleistocene sites, especially in the case of large carnivores.From this perspective, we present here the case of the anthropo-genic utilization of a lion from level TD10-1 of Gran Dolina (Sierrade Atapuerca, Burgos, Spain). The bone remains belonging to Pan-thera leo fossilis show signs of direct interaction between humangroups and big cats from these chronologies. From this perspective,the aim of this paper is to provide data on human use of thesepredators in the European Middle Pleistocene.

2. Methodology

Level TD10-1 faunal analysis of the Gran Dolina site was carriedout following standard archaeozoological methods (Lyman, 1994;Reitz and Wing, 1999) and includes all fossil material from 2000 to2001 excavations. Anatomical, taxonomic and modification detailswere recorded. To assess completeness of the sample, NISP (Numberof Identified Specimens),MNE (MinimumNumber of Elements), MNI(Minimum Number of Individuals) and skeletal survival rate (Brain,1981; Lyman, 1994) were calculated. Skeletal survival rate esti-mates the proportion between the elements recovered and thoseexpected (Brain, 1969). For the calculation of this, the followingformula was used: %survivali¼MNE� 100/number of elementi inthe animal skeleton (MNEe)�MNI.

The age at death of the animals is established mainly from teeth(Silver, 1969; Bökonyi, 1972; Riglet, 1977; Mariezkurrena and Altuna,1983; Azorit et al., 2002). Nevertheless, the epiphyseal fusion and thecortical tissue are also used in this study (Silver, 1969; Barone, 1976).

The breaks on faunal remains were analysed and classifiedaccording to Villa and Mahieu (1991). The outline (transverse,curved/V-shaped, longitudinal), fracture angle (oblique, right,mixed) and fracture edge (smooth, jagged) were analysed.

Alterations observed on bone surfaces were treated at bothmacroscopic and microscopic level. For the microscopic study anOlympus SZ11 stereoscopic (magnification up to 110) and ESEM (FEIQUANTA 600) were used. Damage observed on the faunal remainsincluded cutmarks, intentional bone breakage and carnivoretoothmarks.

The incisions are the only type of cutmark identified. The chop-marks and scrapes are not documented (Binford, 1981; Potts andShipman, 1981; Shipman, 1983; Shipman and Rose, 1983; Shipmanet al., 1984). The incisions are striations with a linear outline ofvariable length, width, and depth. The incisions have a V-shapedsection and display internal microstriation (Potts and Shipman,1981). In some cases, Hertzian cones (Bromage and Boyde, 1984),shoulder effects and barbs (Shipman and Rose,1983) were observed.The analysis of cutmarks took into account the number of striations,location on the anatomical element, distribution over the surface(isolated, clustered, crossed), orientation regarding the longitudinalaxis of the bone (oblique, longitudinal, transverse) and delineation(straight or curved).

Surface damage caused during the bone breakage was also ana-lysed and the diagnostic elements of anthropogenic breakage weredocumented on faunal remains from the TD10-1 sample: percussionpits or percussion marks (Blumenschine and Selvaggio, 1988;Pickering and Egeland, 2006), percussion notches and impactflakes (Capaldo and Blumenschine, 1994; Pickering and Egeland,2006). Percussion marks refer to pits and striae. Diagnosticmorphology and configuration of percussion pits and striations aredescribed by Blumenschine and Selvaggio (1988). Pits are oftenclosely associated with and/or have emanating from them thepatches of striae that result from slippage of stone against bone


during impact events. Percussion notches are semicircular shapedindentations on fracture edges with corresponding negative flakescars. Impact flakes refer to shaft fragments produced by hammer-stone percussion that display the same basic technical attributes ofpercussion as it occurs on stone flakes.

The carnivore toothmarks identified on bone remains in thesample of TD10-1 were mainly pits, punctures and scores (Haynes,1980, 1983b; Binford, 1981; Stiner, 1994; Blumenschine, 1995; interalia). Pits consist of superficial marks lacking a long axis, andpunctures are deep pits that penetrate deeply through the corticalbone or along the edges of an intact or broken bone. Scores aresurface marks with a longitudinal axis more than four times theperpendicular axis. Regarding the breakage, crenulated edges aremainly observed. These present small negative scars on the fractureand longitudinal breakages.

3. Level TD10-1 at the Gran Dolina site

The Gran Dolina site is one of the many caves located in thekarstic complex of the Sierra de Atapuerca (Burgos). It is a cave about18 m high, filled with sediments of the Lower and Middle Pleisto-cene. These fillings are divided into 11 stratigraphical levels. LevelTD10-1 consists of a red sandy clay matrix with angular limestonecentimeter sized clasts (Parés and Pérez-González,1999). This level isdated by TL, ESR and U-series between 250 and 350 Ky (Falguèreset al., 1999; Berger et al., 2008) (Fig. 1). Lithic artefacts recovered atTD10-1 are classified at a technological level as a transitionalmoment between Mode 2 and Mode 3 and are developed primarilywith localmaterials: neogene flint is themost common rawmaterial.Other used raw materials are quartzite, calcarenite and to a lesserextent, cretaceous flint and quartz. All categories of the operationalchain are represented; however, themost abundant elements are theun-retouched flakes (Rodríguez Álvarez, 2004; Menéndez, 2009).

The sample from TD10-1 analysed here provides 16 species ofmacromammals: Ursus arctos, Canis lupus, Vulpes vulpes, P. leo fossilis,Lynx sp., Hystrix sp., Stephanorhinus cf. hemitoechus, Equus ferus,Equus hydruntinus, Sus scrofa, Megaloceros giganteus, Dama damaclactoniana, Cervus elaphus, Bison sp., Hemitragus bonali, Capreoluscapreolus (Table 1). In general, adult specimens dominate the


Fig. 2. (a) Stratigraphical distribution of the faunal remains from TD10-1 according EW section (X¼ 1051e1075); (b) stratigraphical distribution of the faunal remains from TD10-1according NS section (Y¼ 1000e1024); (c) horizontal distribution of the faunal remains from TD10-1 of Gran Dolina. To avoid problems derived of the layer slope, only the faunalremains contained in a band of 24 cm wide have been projected at vertical level.


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Table 2NISP, MNE, MNI and Skeleton survival rate (%Surv) from TD10-1 lion remains of GranDolina.

Panthera leo fossilis (MNI¼ 1) NISP MNE MNEea %Surv.

Maxillae 1 1 2 50Mandible 2 2 2 100Vertebrae 49Ribs 1 1 13 7.69Scapula 2Pelvic girdle 1Humerus 2Radius 2 1 2 50Ulna 2 1 2 50Carpals 16Metacarpus 2 2 10 20Femur 2Patella 2Tibia 2Fibula 1 1 2 50Calcaneus 1 1 2 50Astragalus 2Tarsals 12Metatarsus 1 1 10 10Phalanges 4 4 54 7.41Total 17 15 189 7.93

a Minimal number of expected elements.


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assemblage (66.67%). The skeletal representation of herbivores ischaracterised by the abundance of stylopodials (femur andhumerus), zygopodials (radius and tibia) andmandibles and by a lowrepresentation of the axial skeleton (vertebrae and ribs). Thisskeletal representation coincides with the anatomical elements withhigh medullary value (Binford, 1981; Emerson, 1993).

Analysis of bone breakage shows that curved/V-shapedpredominate along with oblique angles, and smooth edges. Thisdegree of bone breakage at TD10-1 is related to green bonebreakage according to the criteria established by Villa and Mahieu(1991). Surface damage caused during breakage of the bones hasalso been analysed. Diagnostic elements of intentional anthropo-genic breakage are recognised on 1329 bone fragments. Thesediagnostic elements are mainly percussion notches on the shafts oflong bones and impact flakes.

The presence of cutmarks also suggests an association betweenthe hominids and faunal record from TD10-1. Cutmarks are docu-mented on 584 bone remains. These are mainly located on remainsof large andmedium-sized animals (greater than 100 kg inweight).Several cutmarks are associated with the defleshing of largemusclemasses. In these cases, the incisions are long and are locatedlongitudinally or obliquely on the diaphyses of long bones. Shortand deep cutmarks, related to dismemberment and disarticulationof the anatomical portions, are also identified on some epiphyses.

On the other hand, carnivore damage (toothmarks and bonebreakage) on the faunal assemblage is low. Only 4.09% of the boneremains show carnivore activities. These marks are documented onall the animals, especially on medium and small-size animals(individuals with weight below 100 kg). Pits, scores and crenulatededges are the most abundant marks. These are mainly located ondiaphysis fragments. These characteristics do not coincide withthose described for the carnivore dens (Cruz-Uribe, 1991).

Several types of evidences provide information about themethods used to obtain animal nutrients. Some researchers suggestthat themain element to be considered is the skeletal representationof the different taxa identified in the assemblage (Binford, 1981;Brain, 1981; Bunn, 1981; Shipman and Rose, 1983; Klein, 1989;Lyman, 1994; Stiner, 1994; Marean and Kim, 1998; inter alia). Thismust be considered in conjunction with the age profiles of theanimals (Gaudzinski and Roebroeks, 2000) and the localisation ofprocessing marks on the carcasses (Domínguez-Rodrigo, 1999;Domínguez-Rodrigo and Rayne Pickering, 2003; Domínguez-Rodrigo and Barba, 2006; inter alia). From this perspective, thecharacteristics identifiedat TD10-1 faunal assemblage (adult animals,skeletal elements with high nutritional value, cutmarks related tolarge muscle masses removal and low impact of carnivores on thefaunal assemblage) suggest that the anthropic access to animals ismainly primary and immediate, so, the TD10-1 hominids obtain theprey mainly from hunting practices.

Similar characteristics are identified in the work of Rosell(2001), who studied the 1998e1999 excavation seasons fromTD10. From this perspective, continuity can be observed on faunalrecords from upper sublevels of TD10.

4. Data presentation

The sample from TD10-1 (excavation seasons 2000e2001)provided 11,081 faunal remains, of which 1811 were identifiedtaxonomically. Among bones identified at the species level, 47(2.59%) belong to carnivores: U. arctos, C. lupus, V. vulpes, P. leo fossilisand Lynx sp. Of these, 17 remains are attributed to P. leo fossilis.

The archaeostratigraphical distribution of lion fossils in thefaunal assemblage shows a line clearly visible in both the NS sectionas the EW. Therefore, we can distinguish at least a single eventrelated to lions (Fig. 2a, b).


The MNE is 15, mostly represented by phalanges (4), metacarpi(2) and mandibles (2) while the rest of the anatomical elements areonly represented by one bone. Two refits are located in the assem-blage: two fragments from the same radius and two fragments fromthe same ulna. The MNI is 1 established from the most commonskeletal element (mandible) comparing paired elements, age andsize. Regarding age of death, the identified individual is adult.

According to the skeletal survival rate, a biased anatomicalrepresentation is observed (Table 2). The most striking thing is thecomplete absence of vertebrae, pelvic girdles, tibiae and femuri. Wemust bear in mind that the high degree of fragmentation observedin the bone assemblage often makes both specific and anatomicalidentification difficult. From this perspective, it is possible that theseabsent elements, especially those belonging to limb bones, areincluded in the category of long bones ofmedium-sized animals andhave not been identified at the highest level. Because of the highdispersion observed on lion remains, it is also possible that thesemissing elements are in an area not excavated. The TD10-1 exca-vation, despite its wide extension, presents unexcavated areas, suchas east and south areas of the site where stratigraphic profiles havebeen left and the west area, where the railway trench made at theturn of the 19the20th century has disrupted the sediments (Fig. 2c).

On the other hand, the faunal assemblage is dominated by freshbone fragmentation. Analysis of bone breakage shows that curved/V-shapes predominate, along with oblique angles and smoothedges (Villa and Mahieu, 1991). Nevertheless, post-depositionalfragmentation of bones is also identified, specifically the frag-mentation by pressure of sediments. Therefore, stepped orcolumnar fractures are also observed (Marshall, 1989; Lyman,1994). In the case of lion remains, both types of fragmentationare observed: fresh on a radius and dry on an ulna (Fig. 3).

Most of the alterations observed on the lion remains are bothanthropogenic (17.64%) and made by carnivores (29.41%). Therelatively high occurrence of cutmarks on lion bones (11.76%)indicates an association between hominids and this predator. Theseare identified on a phalanx II and on a rib. Incisions on ventralsurface of the rib are predominant (Table 3) (Fig. 4). The actionperformed (viscera removal, skinning and defleshing) is recordedaccording to morphology, emplacement and distribution ofanthropogenic incisions. Surface damage caused during breakage of


Fig. 3. Bone breakage on Panthera leo fossilis remains from TD10-1 of Gran Dolina: (a) fresh bone fragmentation on lion radius; (b) dry bone fragmentation (sediment pressure) onlion ulna.


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the bones is also analysed. One percussion notch is located ona proximal metaphysis of a radius.

As regards carnivore damage, 29.41% of lion remains showmodifications in form of pits and scores (Table 4). The dimensionsand morphologies of the toothmarks mainly coincide with small-sized carnivores, such as foxes (Andrews, 1995; Andrews andFernández-Jalvo, 1997; Selvaggio and Wilder, 2001; Domínguez-Rodrigo and Piqueras, 2003; Yravedra, 2003e04).

On the other hand, 30 remains belonging to other carnivoreswere recovered at the same level. These correspond to 6 individuals:2 wolf, 2 foxes, 1 lynx and 1 bear. All the animals are adults except2 infantile belonging to C. lupus and V. vulpes. The frequency ofmodifications caused by hominids and carnivores on these remainsis low: 1 bone fragment presents cutmarks (3.33%) and 6 showcarnivore damage (20%). The anthropogenic activity is only docu-mented on the pelvic girdle of V. vulpes adult individual, specificallyone cutmark (oblique incision) on the lateral surface of the isquium.On the other hand, toothmarks (mainly pits) are observed ona metatarsus IV of C. lupus, and a mandible and a calcaneus of V.vulpes. Furthermore, a diaphyseal cylinder on a radius of fox and twodigested teeth of U. arctos are identified. However, the majority ofthe carnivore remains are not modified (76.67%) (Table 5). Excludingthe cutmarks documented on an adult fox and on an adult lion, thepresence of these non-human predators in this faunal assemblagemay suggest sporadic events of small and large carnivores during

Table 3Number of Panthera leo fossilis bones with cutmarks at TD10-1 sample of Gran Dolina(obl: oblique; long: longitudinal and tr: transverse).

Rib Phalanx II

No. of Bone remains 1 1Type Incisions IncisionsNo. of striation by

group (min/max)1e16 3e5

Distribution Clustered ClusteredLocation Diaphysis Distal metaphysis

Bone surface Ventral surface AnteriorExternal surface Lateral

Orientation obl oblDelineation Straight Straight

Curved

Measures (mm) 4.1e2.7 1.9e1.66.2e3.1

Action performed Viscera removal Skinning ortendum removalDefleshing


the TD10-1 formation at moments when is was inhabited by humangroups.

5. Discussion

The lion remains identified in the TD10-1 sample of Gran Dolinashow sufficient evidence to attribute their presence to humanactivity during the Middle Pleistocene, specifically MIS 9. Thepresence of cutmarks on bones indicates an association betweenthe hominids and the lion. These marks have enabled us toreconstruct the human consumption sequence of this predator.However, ascertaining the technique of obtaining these largecarnivores is a complex matter. In ethnoarchaeological terms, somecases of lion hunting are present, such as those documented inKenya and Tanzania by the Maasai, in Namibia by the Khoikhois(Hottentots) or in the North of Tanzania by the Hadza. In the case ofthe Maasai, the hunting of lions is related to initiation rites inwhich, after killing the predator, the initiated obtains the group’srespect (Saitori and Beckwith, 1980; Saitoti, 1988). A different caseis the one observed among the Hottentots. In the past, this grouppracticed lion hunting with dogs (African Lion Hound) with the aimof eliminating competitors and protecting their livestock (Lutman,1993). On the other hand, the Hadza kill large carnivores whena confrontation is produced when obtaining prey. O’Connell et al.(1988) documented an episode of lion hunting as a result ofa direct confrontation for the prey.

In the case of TD10-1, the cutmarks identified on the ventralsurface of a lion rib indicate anthropic access to the viscera. Theinternal organs of the thorax are the first parts to disappear in theconsumption sequence of hunting carnivores (Binford, 1981;Blumenschine, 1985, 1986a,b; O’Connell et al., 1988; Selvaggio,1994; Domínguez-Rodrigo, 1994; Capaldo, 1997; Domínguez-Rodrigo, 1999). For this reason, if the TD10-1 hominids eviscer-ated, the access to this predator should be primary and immediateand the hunting could have been the technique used to obtain it.

The purpose of the lion capture is difficult to show archaeo-logically. Besides, this is an isolated case in a context, in which themain prey are red deer and horses and, therefore, the hunting ofa big cat should be understood more as an opportunistic andsporadic episode, rather than a regular activity of the humangroups at this site. Despite being an isolated case, the lion huntingsuggests that Middle Pleistocene hominids are capable of facingthis type of animals successfully. We must take into account thatthe lions are situated in a very high position within the food chainand obtaining them is dangerous and fraught with risk. The factthat no pathologies have been documented on the P. leo fossilis


Fig. 4. Stereoscopic view (binocular SZ11) of cutmarks on Panthera leo fossilis remains identified at TD10-1 sample of Gran Dolina: (a, c, e) incisions on dorsal surface of a lion rib;(b) incisions on ventral surface of lion rib; (d, f) incisions on distal metaphysis of a lion second phalanx.


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remains, which indicate possible diseases or injuries of a traumaticnature that make this predator vulnerable, suggests that thehominids could be located in a similar position in the food chain oreven higher up than these large carnivores.

Table 4Number of Panthera leo fossilis bones with carnivore toothmarks at TD10-1 sample of Gr

No. of boneremains

No. of carnivoredamage

Type Distrib

Rib 1 2 Pits ClusterUlna 1 5 Pit score ClusterMetacarpus II 1 2 Scores Cluster

Metatarsus III 1 1 Pit IsolatePhalanx II 1 2 Pits Disper

Total 5 (29.41%)


Once the lion has been obtained, the processing begins, startingwith the skinning and the viscera removal. Cutmarks located on thedistal metaphysis of a second phalanx can be related to the skinningor tendum removal. The skin and tendum are potentially resources

an Dolina.

ution Location Bone surface Measures (mm)

ed Diaphysis Lateral 3.7� 1.9; 2.3� 1.7ed Diaphysis Medial 4.1� 0.9; 2.1� 1.1; 1.6ed Diaphysis Lateral 0.8� 0.6; 0.3� 1.2

Distal metaphysisd Diaphysis Posterior 2.8� 1.4sed Diaphysis Anterior 1.8� 1.5; 2.5� 2.4

Distal metaphysis


Table 5NISP, MNE, MNI by ages and carnivore and human damage on carnivore remains(excepting the lion) recovered in faunal sample from TD10-1.

NISP MNE Anthropogenicdamage

Carnivoredamage

MNI

ad. inf.

Canislupus

Maxillae 2 2 1 1Mandible 5 1Ulna 2 2Metatarsus IV 1 1 Toothmarks

(pits)

Vulpesvulpes

Mandible 6 3 Toothmarks(pits)

1 1

Scapula 1 1Pelvic girdle 1 1 CutmarksRadius 2 2 Diaphyseal

cylinderMetacarpus III 2 2Metatarsus III 1 1Phalax I 1 1Calcaneus 2 2 Toothmarks

(pits)

Lynx sp. Scapula 1 1 1

Ursusarctos

Mandible 2 1 Two digestedteeth

1

Fibulae 1 1

Total 30 22 1 7 4 2


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for human groups and these can have different functions. In any case,the skinning is required to access the meat and then the content ofthe bones. Cutmarks related to defleshing are also documented onexternal surface of a rib. These incisions are associated with the backmuscles removal (Trapezius and/or Latissimus dorsi).

With the available elements, two fracturing techniques can besuggested in the case of the radius: (1) active or thrown percussionand (2) passive or direct percussion (Giusberti and Peretto, 1991;Anconetani, 1999). Active or thrown percussion is produced whenthe skeletal element is hit directly against an object (stone) andpassive or direct percussion is caused when the bone is held on theground or on an object that acts as an anvil and is hit with aninstrument generally made of stone (hammerstone).

Once the resources have been used, the lion bones are leftbehind like the rest of the animals processed at TD10-1. Once thehuman groups leave the camp, the smell given off by the remainscan attract other predators that act as scavengers in search ofpotentially consumable resources (Binford, 1981; Rosell and Blasco,2008, 2009). The number and dimensions of the toothmarksidentified on the lion skeleton may be due to the activity of smallscavengers, probably small canines, such as foxes. Nevertheless, atcertain times, toothmarks of larger dimensions have also beenobserved, which could be related to large predators, such as hyenas.Further evidence, which corroborates the marauding activity of thecarnivores at TD10-1, is the presence of overlapping marks on someherbivores remains (Blasco and Rosell, 2009). These correspond tocarnivore toothmarks situated over cutmarks.

In general, this study provides data on the relationship betweenhuman groups and big cats of the Middle Pleistocene from evidencebasedon the occasional anthropic use of a lion fromTD10-1. Althoughthe utilization of carnivores is not isolated at this level (cutmarks onisquium of an adult fox), the processingmarks on big cat remains arenot common at sites of similar chronologies. However, in moremodern chronologies, the specific use of other large carnivores, suchas the cave bear (Ursus spelaeus), has been suggested at some sites ofthe European Middle Palaeolithic. This is the case of Arcy-sur-Cure(David, 1997), Biache Saint Vast (Auguste, 1991, 1995) and Prélétang(Tillet, 2002) in France or of Sant’Agostino (Stiner, 1994), the Grottadelle Fate and Madonna dell’Arma in Italy (Tillet, 2002). Another


example is the cutmarks located on a dhole (Cuon cf. alpinus)mandible from Cova Negra (Pérez Ripoll et al., 2010) or the use of theskin of a leopard (Panthera pardus) at the site of Los Torrejones Cave(Arribas et al., 1997), both in Spain. Nevertheless, most of the exam-ples of anthropic use of carnivores come from Upper Palaeolithicsites. From this perspective, there are numerous cases of the utili-zation of the lynx for food in some localities such as Peña de laEstebanvela (Yravedra, 2005), Cova Beneito or Les Cendres (MartínezValle, 1996) in Spain and some examples of the anthropic processingof the badger, fox and bear at the Italian site of Polesini (Stiner,1994).Evidence has also been documented on fox remains at some Frenchsites, such as Combe Saunière (Chauvière and Castel, 2004).

6. Conclusions

The study of the relationship between hominids and carnivoresduring the Pleistocene has currently been a highly debated topic.Relations between both predators have been interpreted inmultipleways: lack of contact between them, relations of dependency orcompetition for resources and occupied space, etc. The anthropo-genic use of these animals as food is not common at the MiddlePleistocene sites, especially in the case of big cats. From thisperspective, the TD10-1 sample provides sufficient evidence toshow the human use of a lion (P. leo fossilis) during MIS 9. Thediagnostic elements of human activity, identified on the lionremains, allow us to document some episodes of the processingsequence: cutmarks related to the skinning and defleshing areidentified and the human use of bone marrow is documented bydiagnostic elements of anthropogenic breakage. All these evidencessuggest that the lion was used for food.

On the other hand, cutmarks identifiedonventral surface on a riband related to viscera removal can only be understood froma primary and immediate anthropic access to the animal. Therefore,the hunting may be the most probable technique used by thehominids to catch the lion. From this, the direct interaction betweenhominids and this big cat at level TD10-1 is documented. However,this is an isolated case in a context, in which the main prey areungulates, and therefore, the hunting of a lion should be understoodas a sporadic episode rather than a systematic activity of the humangroups at this site. Nevertheless, the hunting of this predatorsuggests that the hominids of the Middle Pleistocene are successfulhunters able to face the large predators. In this paper, we haveshown that the hunting of large carnivores is another possibility forunderstanding the relationships between hominids and carnivoresduring the Middle Pleistocene.

Acknowledgements

Thanks to all the members of the Atapuerca research team,especially the one from the Gran Dolina site. Special thanks to HelleKettner for English corrections. This research was supported by theMinisterio de Educación y Ciencia Spanish Government GrantsCGL2009-12703-C03-01, CGL2009-12703-C03-02, CGL2009-12703-C03-03 and CGL2009-7896, and by the Generalitat de CatalunyaGrant 2009 SGR 188. Ruth Blasco is beneficiary of a FI Grant from theGeneralitat de Catalunya and financed by the European Social Found.The field excavation work was supported by the Junta de Castilla yLeón and Fundación Atapuerca.

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