(Panthera Leo Fossilis) at the Gran Dolina Site, Sierra De Atapuerca, Spain

Journal of Archaeological Science 37 (2010) 2051e2060

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Journal of Archaeological Science

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The hunted hunter: the capture of a lion (Panthera leo fossilis) at the Gran 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, Spain b Departamento de Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain c Centro de Investigación (UCM-ISCIII) de Evolución y Comportamiento Humanos, C/Sinesio Delgado, 4 (Pabellón 14), 28029 Madrid, Spain d CENIEH (Centro Nacional de Investigación sobre Evolución Humana), Avenida de la Paz 28, 09004 Burgos, Spain e Visiting professor, Institute of Vertebrate Paleontology and Paleoanthropology of Beijing (IVPP) article info abstract

Article history: Many Pleistocene caves and rock shelters contain evidence of carnivore and human activities. For this Received 22 December 2009 reason, it is common to recover at these sites faunal remains left by both biological agents. In order to Received in revised form explain the role that carnivores play at the archaeological sites it is necessary to analyse several elements, 15 March 2010 such as the taxonomical and skeletal representation, the age proﬁles, the ratio of NISP to MNI, the Accepted 17 March 2010 anthropogenic processing marks on the carcasses (location and purpose of cutmarks and burning and bone breakage patterns), carnivore damage (digested bones, location and frequencies of toothmarks and bone Keywords: breakage), length of the long bones, frequencies of coprolites and vertical distribution of the faunal Subsistence strategies Hunting remains, inter alia. From this, the documentation of carnivores in a faunal assemblage with a clear Carnivore use anthropogenic component can be understood from three main phenomena: (1) the carnivores as accu- Panthera leo fossilis mulators and the use of the site as a den; (2) carnivores as scavengers of hominid refuse and; (3) carnivores Gran Dolina as hominids’ prey. Of these three phenomena, the last one is the least documented at the Middle Pleis- Sierra de Atapuerca tocene sites. From this perspective, here we present the case of the anthropogenic use of a lion (Panthera Middle Pleistocene leo fossilis) from level TD10-1 of Gran Dolina (MIS 9, Sierra de Atapuerca, Burgos, Spain). The lion bone remains show signs of direct interaction between this big cat and human groups that occupied Gran Dolina in these chronologies. From this perspective, the aim of this paper is to contribute to the knowledge of the role developed by large carnivores in the anthropogenic contexts and to provide data on human use of these predators at the European Middle Pleistocene sites. Ó 2010 Elsevier Ltd. All rights reserved.

1. Introduction alia). However, we must bear in mind that not all carnivores behave in a similar way or have the same impact on the carcasses. Each species The archaeological excavations carried out at the Middle Pleisto- has its own ethology and physical characteristics, which influence the cene sites show the occupation of karstic areas by hominids and accumulations that they produce and the intensity with which they carnivores. For this reason, the recovery of faunal remains left by both act on bone remains. Even so, a series of general characteristics are biological entities is common at these sites (both the bones left identified in all assemblage generated by non-human predators. In behind from their activities and/or their own skeletal remains). Many general, the age profiles of the prey tend to be attritional (with a carnivores regularly use caves as a breeding place or as a refuge. There predominance of young and senile individuals over adults). The are several studies that allow us to recognise both current and skeletal representation of the prey is highly variable with a domi- archaeological dens (Sutcliffe, 1970; Haynes, 1980, 1983a; Hill, 1984; nance of limb bones (except for carpal and tarsal bones which can be Blumenschine, 1985, 1986a, 1988a, 1988b; Clot, 1987; Cruz-Uribe, swallowed during the consumption process) and some elements of 1991; Domínguez-Rodrigo, 1994, 2001; Selvaggio, 1994; Fosse, the axial and cranial skeleton (Binford, 1981). The proportion 1996; Villa and Soressi, 2000; Villa et al., 2004; Michel, 2005; inter between the cranialepostcranial skeleton of the prey decreases according to the accumulating carnivore and to the size of the ungulates, so the larger the prey, the lower the capacity of the carnivore to transport the skull (Cruz-Uribe, 1991). Furthermore, * Corresponding author. Tel.: þ34 977 257 882; fax: þ34 977559597. E-mail address: [email protected] (R. Blasco). coprolites, skeletal remains belonging to other carnivores, deciduous

0305-4403/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.jas.2010.03.010 2052 R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060 teeth (in the case of hyena maternity dens) and digested bones are elements of the axial skeleton. The modifications on these often retrieved at these assemblages (Cruz-Uribe, 1991; Villa et al., anatomical portions are so intense that in many cases they make 2004; inter alia). Some authors, such as Fosse (1994) or Villa et al. them disappear. On the contrary, the diaphyses, which are generally (2004), also observed differences related to the bone fragmentation highly fractured during the processing and anthropic consumption, between the carnivore sites and anthropogenic sites. According to show very little impact. This duality is due to the fact that bones these authors, the bones recovered in the assemblages generated by with a high content of spongy tissue (epiphyses and flat bones) carnivores usually are less fragmented and, therefore, the propor- contain resources (fat), which are difficult to reach for the hominids tions of identifiable elements are much higher. without the appropriate technology, such as boiling (Oliver, 1993). However, the carnivores are not only accumulators. The smells However, during this process, carnivores do not only destroy the from the remains left in the human camps are attractive for many bones, but they can also affect the original position of the remains, predators and, therefore, it is common for these carnivores to act as by carrying out important remobilisations and significantly altering scavengers in search of potentially consumable elements (Binford, the spatial distribution left by human groups (Binford et al., 1988). 1981). Different observations and experimental reproductions Nevertheless, the presence of carnivores and their damage in have been made with wild and captive animals, attempting to the anthropogenic context, do not only respond to the scavenging document the carnivore damage on the faunal assemblages gener- activities or to the use of the site as den. Carnivores can also be ated by human groups (Sutcliffe, 1970; Klein, 1975; Bunn, 1986; a potential prey for human groups because they can offer a series of Bunn et al., 1980; Bunn et al., 1988; Blumenschine, 1986a, 1986b; usable resources (skin, tendons, meat, marrow, etc.). As with Marean et al., 1992). All of these studies coincide with the fact that the herbivores, the processing of the carnivores is identified by the scavengers have a predilection for epiphyses of limb bones and for presence of cutmarks, the intentional bone breakage and/or the

Fig. 1. Location, composite stratigraphic profile of deposits and levels dating at the Gran Dolina site from Parés and Pérez-González (1999), Falguères et al. (1999) and Berger et al. (2008). R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060 2053 burning patterns of their remains. Although archaeological exam- during impact events. Percussion notches are semicircular shaped ples exist in some assemblages of the first half of the Upper Pleis- indentations on fracture edges with corresponding negative flake tocene (Auguste, 1991,1995; Stiner, 1994; Arribas et al., 1997; David, scars. Impact flakes refer to shaft fragments produced by hammer- 1997; Tillet, 2002), the use of these animals is not common at the stone percussion that display the same basic technical attributes of Middle Pleistocene sites, especially in the case of large carnivores. percussion as it occurs on stone flakes. From this perspective, we present here the case of the anthropo- The carnivore toothmarks identified on bone remains in the genic utilization of a lion from level TD10-1 of Gran Dolina (Sierra sample of TD10-1 were mainly pits, punctures and scores (Haynes, de Atapuerca, Burgos, Spain). The bone remains belonging to Pan- 1980, 1983b; Binford, 1981; Stiner, 1994; Blumenschine, 1995; inter thera leo fossilis show signs of direct interaction between human alia). Pits consist of superficial marks lacking a long axis, and groups and big cats from these chronologies. From this perspective, punctures are deep pits that penetrate deeply through the cortical the aim of this paper is to provide data on human use of these bone or along the edges of an intact or broken bone. Scores are predators in the European Middle Pleistocene. surface marks with a longitudinal axis more than four times the perpendicular axis. Regarding the breakage, crenulated edges are 2. Methodology mainly observed. These present small negative scars on the fracture and longitudinal breakages. Level TD10-1 faunal analysis of the Gran Dolina site was carried out following standard archaeozoological methods (Lyman, 1994; 3. Level TD10-1 at the Gran Dolina site Reitz and Wing, 1999) and includes all fossil material from 2000 to fi 2001 excavations. Anatomical, taxonomic and modi cation details The Gran Dolina site is one of the many caves located in the were recorded. To assess completeness of the sample, NISP (Number karstic complex of the Sierra de Atapuerca (Burgos). It is a cave about fi of Identi ed Specimens), MNE (Minimum Number of Elements), MNI 18 m high, filled with sediments of the Lower and Middle Pleisto- (Minimum Number of Individuals) and skeletal survival rate (Brain, cene. These fillings are divided into 11 stratigraphical levels. Level 1981; Lyman, 1994) were calculated. Skeletal survival rate esti- TD10-1 consists of a red sandy clay matrix with angular limestone mates the proportion between the elements recovered and those centimeter sized clasts (Parés and Pérez-González,1999). This level is expected (Brain, 1969). For the calculation of this, the following dated by TL, ESR and U-series between 250 and 350 Ky (Falguères ¼ formula was used: %survivali MNE 100/number of elementi in et al., 1999; Berger et al., 2008)(Fig. 1). Lithic artefacts recovered at the animal skeleton (MNEe) MNI. TD10-1 are classified at a technological level as a transitional The age at death of the animals is established mainly from teeth moment between Mode 2 and Mode 3 and are developed primarily (Silver, 1969; Bökonyi, 1972; Riglet, 1977; Mariezkurrena and Altuna, with local materials: neogene flint is the most common raw material. 1983; Azorit et al., 2002). Nevertheless, the epiphyseal fusion and the Other used raw materials are quartzite, calcarenite and to a lesser cortical tissue are also used in this study (Silver, 1969; Barone, 1976). extent, cretaceous flint and quartz. All categories of the operational fi The breaks on faunal remains were analysed and classi ed chain are represented; however, the most abundant elements are the according to Villa and Mahieu (1991). The outline (transverse, un-retouched flakes (Rodríguez Álvarez, 2004; Menéndez, 2009). curved/V-shaped, longitudinal), fracture angle (oblique, right, The sample from TD10-1 analysed here provides 16 species of mixed) and fracture edge (smooth, jagged) were analysed. macromammals: Ursus arctos, Canis lupus, Vulpes vulpes, P. leo fossilis, Alterations observed on bone surfaces were treated at both Lynx sp., Hystrix sp., Stephanorhinus cf. hemitoechus, Equus ferus, macroscopic and microscopic level. For the microscopic study an Equus hydruntinus, Sus scrofa, Megaloceros giganteus, Dama dama fi Olympus SZ11 stereoscopic (magni cation up to 110) and ESEM (FEI clactoniana, Cervus elaphus, Bison sp., Hemitragus bonali, Capreolus QUANTA 600) were used. Damage observed on the faunal remains capreolus (Table 1). In general, adult specimens dominate the included cutmarks, intentional bone breakage and carnivore toothmarks. The incisions are the only type of cutmark identified. The chop- Table 1 marks and scrapes are not documented (Binford, 1981; Potts and NISP, MNE and MNI by ages from TD10-1 faunal sample of Gran Dolina. Shipman, 1981; Shipman, 1983; Shipman and Rose, 1983; Shipman TD10-1 Taxa NISP MNE MNI MNI by ages et al., 1984). The incisions are striations with a linear outline of neo. inf. juv. ad. sen. variable length, width, and depth. The incisions have a V-shaped Ursus arctos 321 1 section and display internal microstriation (Potts and Shipman, Canis lupus 10 6 2 1 1 1981). In some cases, Hertzian cones (Bromage and Boyde, 1984), Vulpes vulpes 16 13 2 1 1 shoulder effects and barbs (Shipman and Rose, 1983)wereobserved. Panthera leo 17 15 1 1 The analysis of cutmarks took into account the number of striations, Lynx sp. 1 1 1 1 location on the anatomical element, distribution over the surface Hystrix sp. 2 1 1 1 Stephanorhinus 52 9 2 1 1 (isolated, clustered, crossed), orientation regarding the longitudinal cf. hemitoechus axis of the bone (oblique, longitudinal, transverse) and delineation Equus ferus 260 62 9 2 3 3 1 (straight or curved). Equus hydruntinus 12 5 2 1 1 Surface damage caused during the bone breakage was also ana- Sus scrofa 111 1 Cervidae 121 24 2 1 1 lysed and the diagnostic elements of anthropogenic breakage were Megaloceros giganteus 111 1 documented on faunal remains from the TD10-1 sample: percussion Dama dama clactoniana 221 1 pits or percussion marks (Blumenschine and Selvaggio, 1988; Cervus elaphus 762 232 9 1 1 6 1 Pickering and Egeland, 2006), percussion notches and impact Bison sp. 144 55 5 1 1 2 1 flakes (Capaldo and Blumenschine, 1994; Pickering and Egeland, Hemitragus bonali 551 1 Capreolus capreolus 332 1 1 2006). Percussion marks refer to pits and striae. Diagnostic Erinaceidae 11 8 1 1 morphology and configuration of percussion pits and striations are Leporidae 329 167 12 1 11 described by Blumenschine and Selvaggio (1988). Pits are often Aves 58 40 3 3 closely associated with and/or have emanating from them the Ichthyofauna 1 1 1 1 Total 1811 651 60 1 9 7 40 3 patches of striae that result from slippage of stone against bone 2054 R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060

Fig. 2. (a) Stratigraphical distribution of the faunal remains from TD10-1 according EW section (X ¼ 1051e1075); (b) stratigraphical distribution of the faunal remains from TD10-1 according NS section (Y ¼ 1000e1024); (c) horizontal distribution of the faunal remains from TD10-1 of Gran Dolina. To avoid problems derived of the layer slope, only the faunal remains contained in a band of 24 cm wide have been projected at vertical level. R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060 2055 assemblage (66.67%). The skeletal representation of herbivores is The MNE is 15, mostly represented by phalanges (4), metacarpi characterised by the abundance of stylopodials (femur and (2) and mandibles (2) while the rest of the anatomical elements are humerus), zygopodials (radius and tibia) and mandibles and by a low only represented by one bone. Two refits are located in the assem- representation of the axial skeleton (vertebrae and ribs). This blage: two fragments from the same radius and two fragments from skeletal representation coincides with the anatomical elements with the same ulna. The MNI is 1 established from the most common high medullary value (Binford, 1981; Emerson, 1993). skeletal element (mandible) comparing paired elements, age and Analysis of bone breakage shows that curved/V-shaped size. Regarding age of death, the identified individual is adult. predominate along with oblique angles, and smooth edges. This According to the skeletal survival rate, a biased anatomical degree of bone breakage at TD10-1 is related to green bone representation is observed (Table 2). The most striking thing is the breakage according to the criteria established by Villa and Mahieu complete absence of vertebrae, pelvic girdles, tibiae and femuri. We (1991). Surface damage caused during breakage of the bones has must bear in mind that the high degree of fragmentation observed also been analysed. Diagnostic elements of intentional anthropo- in the bone assemblage often makes both specific and anatomical genic breakage are recognised on 1329 bone fragments. These identification difficult. From this perspective, it is possible that these diagnostic elements are mainly percussion notches on the shafts of absent elements, especially those belonging to limb bones, are long bones and impact flakes. included in the category of long bones of medium-sized animals and The presence of cutmarks also suggests an association between have not been identified at the highest level. Because of the high the hominids and faunal record from TD10-1. Cutmarks are docu- dispersion observed on lion remains, it is also possible that these mented on 584 bone remains. These are mainly located on remains missing elements are in an area not excavated. The TD10-1 exca- of large and medium-sized animals (greater than 100 kg in weight). vation, despite its wide extension, presents unexcavated areas, such Several cutmarks are associated with the defleshing of large muscle as east and south areas of the site where stratigraphic profiles have masses. In these cases, the incisions are long and are located been left and the west area, where the railway trench made at the longitudinally or obliquely on the diaphyses of long bones. Short turn of the 19the20th century has disrupted the sediments (Fig. 2c). and deep cutmarks, related to dismemberment and disarticulation On the other hand, the faunal assemblage is dominated by fresh of the anatomical portions, are also identified on some epiphyses. bone fragmentation. Analysis of bone breakage shows that curved/ On the other hand, carnivore damage (toothmarks and bone V-shapes predominate, along with oblique angles and smooth breakage) on the faunal assemblage is low. Only 4.09% of the bone edges (Villa and Mahieu, 1991). Nevertheless, post-depositional remains show carnivore activities. These marks are documented on fragmentation of bones is also identified, specifically the frag- all the animals, especially on medium and small-size animals mentation by pressure of sediments. Therefore, stepped or (individuals with weight below 100 kg). Pits, scores and crenulated columnar fractures are also observed (Marshall, 1989; Lyman, edges are the most abundant marks. These are mainly located on 1994). In the case of lion remains, both types of fragmentation diaphysis fragments. These characteristics do not coincide with are observed: fresh on a radius and dry on an ulna (Fig. 3). those described for the carnivore dens (Cruz-Uribe, 1991). Most of the alterations observed on the lion remains are both Several types of evidences provide information about the anthropogenic (17.64%) and made by carnivores (29.41%). The methods used to obtain animal nutrients. Some researchers suggest relatively high occurrence of cutmarks on lion bones (11.76%) that the main element to be considered is the skeletal representation indicates an association between hominids and this predator. These of the different taxa identified in the assemblage (Binford, 1981; are identified on a phalanx II and on a rib. Incisions on ventral Brain, 1981; Bunn, 1981; Shipman and Rose, 1983; Klein, 1989; surface of the rib are predominant (Table 3)(Fig. 4). The action Lyman, 1994; Stiner, 1994; Marean and Kim, 1998; inter alia). This performed (viscera removal, skinning and defleshing) is recorded must be considered in conjunction with the age profiles of the according to morphology, emplacement and distribution of animals (Gaudzinski and Roebroeks, 2000) and the localisation of anthropogenic incisions. Surface damage caused during breakage of processing marks on the carcasses (Domínguez-Rodrigo, 1999; Domínguez-Rodrigo and Rayne Pickering, 2003; Domínguez-

Rodrigo and Barba, 2006; inter alia). From this perspective, the Table 2 characteristics identified at TD10-1 faunal assemblage (adult animals, NISP, MNE, MNI and Skeleton survival rate (%Surv) from TD10-1 lion remains of Gran skeletal elements with high nutritional value, cutmarks related to Dolina. large muscle masses removal and low impact of carnivores on the Panthera leo fossilis (MNI ¼ 1) NISP MNE MNEea %Surv. faunal assemblage) suggest that the anthropic access to animals is Maxillae 1 1 2 50 mainly primary and immediate, so, the TD10-1 hominids obtain the Mandible 2 2 2 100 prey mainly from hunting practices. Vertebrae 49 Similar characteristics are identified in the work of Rosell Ribs 1 1 13 7.69 (2001), who studied the 1998e1999 excavation seasons from Scapula 2 Pelvic girdle 1 TD10. From this perspective, continuity can be observed on faunal Humerus 2 records from upper sublevels of TD10. Radius 2 1 2 50 Ulna 2 1 2 50 4. Data presentation Carpals 16 Metacarpus 2 2 10 20 Femur 2 e The sample from TD10-1 (excavation seasons 2000 2001) Patella 2 provided 11,081 faunal remains, of which 1811 were identified Tibia 2 taxonomically. Among bones identified at the species level, 47 Fibula 1 1 2 50 (2.59%) belong to carnivores: U. arctos, C. lupus, V. vulpes, P. leo fossilis Calcaneus 1 1 2 50 Astragalus 2 and Lynx sp. Of these, 17 remains are attributed to P. leo fossilis. Tarsals 12 The archaeostratigraphical distribution of lion fossils in the Metatarsus 1 1 10 10 faunal assemblage shows a line clearly visible in both the NS section Phalanges 4 4 54 7.41 as the EW. Therefore, we can distinguish at least a single event Total 17 15 189 7.93 related to lions (Fig. 2a, b). a Minimal number of expected elements. 2056 R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060

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) on lion ulna. the bones is also analysed. One percussion notch is located on the TD10-1 formation at moments when is was inhabited by human a proximal metaphysis of a radius. groups. As regards carnivore damage, 29.41% of lion remains show modifications in form of pits and scores (Table 4). The dimensions 5. Discussion and morphologies of the toothmarks mainly coincide with small- sized carnivores, such as foxes (Andrews, 1995; Andrews and The lion remains identified in the TD10-1 sample of Gran Dolina Fernández-Jalvo, 1997; Selvaggio and Wilder, 2001; Domínguez- show sufficient evidence to attribute their presence to human Rodrigo and Piqueras, 2003; Yravedra, 2003e04). activity during the Middle Pleistocene, specifically MIS 9. The On the other hand, 30 remains belonging to other carnivores presence of cutmarks on bones indicates an association between were recovered at the same level. These correspond to 6 individuals: the hominids and the lion. These marks have enabled us to 2 wolf, 2 foxes, 1 lynx and 1 bear. All the animals are adults except reconstruct the human consumption sequence of this predator. 2 infantile belonging to C. lupus and V. vulpes. The frequency of However, ascertaining the technique of obtaining these large modifications caused by hominids and carnivores on these remains carnivores is a complex matter. In ethnoarchaeological terms, some is low: 1 bone fragment presents cutmarks (3.33%) and 6 show cases of lion hunting are present, such as those documented in carnivore damage (20%). The anthropogenic activity is only docu- Kenya and Tanzania by the Maasai, in Namibia by the Khoikhois mented on the pelvic girdle of V. vulpes adult individual, specifically (Hottentots) or in the North of Tanzania by the Hadza. In the case of one cutmark (oblique incision) on the lateral surface of the isquium. the Maasai, the hunting of lions is related to initiation rites in On the other hand, toothmarks (mainly pits) are observed on which, after killing the predator, the initiated obtains the group’s a metatarsus IV of C. lupus, and a mandible and a calcaneus of V. respect (Saitori and Beckwith, 1980; Saitoti, 1988). A different case vulpes. Furthermore, a diaphyseal cylinder on a radius of fox and two is the one observed among the Hottentots. In the past, this group digested teeth of U. arctos are identified. However, the majority of practiced lion hunting with dogs (African Lion Hound) with the aim the carnivore remains are not modified (76.67%) (Table 5). Excluding of eliminating competitors and protecting their livestock (Lutman, the cutmarks documented on an adult fox and on an adult lion, the 1993). On the other hand, the Hadza kill large carnivores when presence of these non-human predators in this faunal assemblage a confrontation is produced when obtaining prey. O’Connell et al. may suggest sporadic events of small and large carnivores during (1988) documented an episode of lion hunting as a result of a direct confrontation for the prey. In the case of TD10-1, the cutmarks identified on the ventral Table 3 surface of a lion rib indicate anthropic access to the viscera. The Number of Panthera leo fossilis bones with cutmarks at TD10-1 sample of Gran Dolina internal organs of the thorax are the first parts to disappear in the (obl: oblique; long: longitudinal and tr: transverse). consumption sequence of hunting carnivores (Binford, 1981; Rib Phalanx II Blumenschine, 1985, 1986a,b; O’Connell et al., 1988; Selvaggio, No. of Bone remains 1 1 1994; Domínguez-Rodrigo, 1994; Capaldo, 1997; Domínguez- Type Incisions Incisions Rodrigo, 1999). For this reason, if the TD10-1 hominids eviscer- No. of striation by 1e16 3e5 ated, the access to this predator should be primary and immediate group (min/max) and the hunting could have been the technique used to obtain it. Distribution Clustered Clustered fi Location Diaphysis Distal metaphysis The purpose of the lion capture is dif cult to show archaeo- logically. Besides, this is an isolated case in a context, in which the Bone surface Ventral surface Anterior main prey are red deer and horses and, therefore, the hunting of External surface Lateral a big cat should be understood more as an opportunistic and Orientation obl obl sporadic episode, rather than a regular activity of the human Delineation Straight Straight Curved groups at this site. Despite being an isolated case, the lion hunting suggests that Middle Pleistocene hominids are capable of facing e e Measures (mm) 4.1 2.7 1.9 1.6 this type of animals successfully. We must take into account that 6.2e3.1 the lions are situated in a very high position within the food chain Action performed Viscera removal Skinning or and obtaining them is dangerous and fraught with risk. The fact Defleshing tendum removal that no pathologies have been documented on the P. leo fossilis R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060 2057

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. remains, which indicate possible diseases or injuries of a traumatic Once the lion has been obtained, the processing begins, starting nature that make this predator vulnerable, suggests that the with the skinning and the viscera removal. Cutmarks located on the hominids could be located in a similar position in the food chain or distal metaphysis of a second phalanx can be related to the skinning even higher up than these large carnivores. or tendum removal. The skin and tendum are potentially resources

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

No. of bone No. of carnivore Type Distribution Location Bone surface Measures (mm) remains damage Rib 1 2 Pits Clustered Diaphysis Lateral 3.7 1.9; 2.3 1.7 Ulna 1 5 Pit score Clustered Diaphysis Medial 4.1 0.9; 2.1 1.1; 1.6 Metacarpus II 1 2 Scores Clustered Diaphysis Lateral 0.8 0.6; 0.3 1.2 Distal metaphysis Metatarsus III 1 1 Pit Isolated Diaphysis Posterior 2.8 1.4 Phalanx II 1 2 Pits Dispersed Diaphysis Anterior 1.8 1.5; 2.5 2.4 Distal metaphysis Total 5 (29.41%) 2058 R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060

Table 5 example is the cutmarks located on a dhole (Cuon cf. alpinus) NISP, MNE, MNI by ages and carnivore and human damage on carnivore remains mandible from Cova Negra (Pérez Ripoll et al., 2010)ortheuseofthe (excepting the lion) recovered in faunal sample from TD10-1. skin of a leopard (Panthera pardus) at the site of Los Torrejones Cave NISP MNE Anthropogenic Carnivore MNI (Arribas et al., 1997), both in Spain. Nevertheless, most of the exam- damage damage ad. inf. ples of anthropic use of carnivores come from Upper Palaeolithic Canis Maxillae 2 2 1 1 sites. From this perspective, there are numerous cases of the utili- lupus Mandible 5 1 zation of the lynx for food in some localities such as Peña de la Ulna 2 2 Estebanvela (Yravedra, 2005), Cova Beneito or Les Cendres (Martínez Metatarsus IV 1 1 Toothmarks Valle, 1996) in Spain and some examples of the anthropic processing (pits) of the badger, fox and bear at the Italian site of Polesini (Stiner, 1994). Vulpes Mandible 6 3 Toothmarks 11 Evidence has also been documented on fox remains at some French vulpes (pits) sites, such as Combe Saunière (Chauvière and Castel, 2004). Scapula 1 1 Pelvic girdle 1 1 Cutmarks Radius 2 2 Diaphyseal 6. Conclusions cylinder Metacarpus III 2 2 The study of the relationship between hominids and carnivores Metatarsus III 1 1 Phalax I 1 1 during the Pleistocene has currently been a highly debated topic. Calcaneus 2 2 Toothmarks Relations between both predators have been interpreted in multiple (pits) ways: lack of contact between them, relations of dependency or Lynx sp. Scapula 1 1 1 competition for resources and occupied space, etc. The anthropogenic use of these animals as food is not common at the Middle Ursus Mandible 2 1 Two digested 1 arctos teeth Pleistocene sites, especially in the case of big cats. From this Fibulae 1 1 perspective, the TD10-1 sample provides sufficient evidence to show the human use of a lion (P. leo fossilis) during MIS 9. The Total 30 22 1 7 4 2 diagnostic elements of human activity, identified on the lion remains, allow us to document some episodes of the processing for human groups and these can have different functions. In any case, sequence: cutmarks related to the skinning and defleshing are the skinning is required to access the meat and then the content of identified and the human use of bone marrow is documented by the bones. Cutmarks related to defleshing are also documented on diagnostic elements of anthropogenic breakage. All these evidences external surface of a rib. These incisions are associated with the back suggest that the lion was used for food. muscles removal (Trapezius and/or Latissimus dorsi). On the other hand, cutmarks identified on ventral surface on a rib With the available elements, two fracturing techniques can be and related to viscera removal can only be understood from suggested in the case of the radius: (1) active or thrown percussion a primary and immediate anthropic access to the animal. Therefore, and (2) passive or direct percussion (Giusberti and Peretto, 1991; the hunting may be the most probable technique used by the Anconetani, 1999). Active or thrown percussion is produced when hominids to catch the lion. From this, the direct interaction between the skeletal element is hit directly against an object (stone) and hominids and this big cat at level TD10-1 is documented. However, passive or direct percussion is caused when the bone is held on the this is an isolated case in a context, in which the main prey are ground or on an object that acts as an anvil and is hit with an ungulates, and therefore, the hunting of a lion should be understood instrument generally made of stone (hammerstone). as a sporadic episode rather than a systematic activity of the human Once the resources have been used, the lion bones are left groups at this site. Nevertheless, the hunting of this predator behind like the rest of the animals processed at TD10-1. Once the suggests that the hominids of the Middle Pleistocene are successful human groups leave the camp, the smell given off by the remains hunters able to face the large predators. In this paper, we have can attract other predators that act as scavengers in search of shown that the hunting of large carnivores is another possibility for potentially consumable resources (Binford, 1981; Rosell and Blasco, understanding the relationships between hominids and carnivores 2008, 2009). The number and dimensions of the toothmarks during the Middle Pleistocene. identified on the lion skeleton may be due to the activity of small scavengers, probably small canines, such as foxes. Nevertheless, at Acknowledgements certain times, toothmarks of larger dimensions have also been observed, which could be related to large predators, such as hyenas. Thanks to all the members of the Atapuerca research team, Further evidence, which corroborates the marauding activity of the especially the one from the Gran Dolina site. Special thanks to Helle carnivores at TD10-1, is the presence of overlapping marks on some Kettner for English corrections. This research was supported by the herbivores remains (Blasco and Rosell, 2009). These correspond to Ministerio de Educación y Ciencia Spanish Government Grants carnivore toothmarks situated over cutmarks. CGL2009-12703-C03-01, CGL2009-12703-C03-02, CGL2009-12703- In general, this study provides data on the relationship between C03-03 and CGL2009-7896, and by the Generalitat de Catalunya human groups and big cats of the Middle Pleistocene from evidence Grant 2009 SGR 188. Ruth Blasco is beneficiary of a FI Grant from the based on the occasional anthropic use of a lion from TD10-1. Although Generalitat de Catalunya and financed by the European Social Found. the utilization of carnivores is not isolated at this level (cutmarks on The field excavation work was supported by the Junta de Castilla y isquium of an adult fox), the processing marks on big cat remains are León and Fundación Atapuerca. not common at sites of similar chronologies. However, in more fi modern chronologies, the speci c use of other large carnivores, such References as the cave bear (Ursus spelaeus), has been suggested at some sites of the European Middle Palaeolithic. This is the case of Arcy-sur-Cure Anconetani, P., 1999. L’assemblage faunique du gisement paléolithique inférieur (David, 1997), Biache Saint Vast (Auguste, 1991, 1995) and Prélétang d’Isernia-La Pineta (Molise, Italie) et l’explotation du bison. In: Brugal, J.P., ’ David, F., Enloe, J.G., Jaubert, J. (Eds.), Le Bison: Gibier et Moyen de Subsistance (Tillet, 2002)inFranceorofSantAgostino (Stiner, 1994), the Grotta des Hommes du Paléolithique aux Paléoindiens des Grandes Plaines. Actes du delle Fate and Madonna dell’Arma in Italy (Tillet, 2002). Another Colloque International, Toulouse, 1995. APDCA, Antibes, pp. 105e120. R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060 2059

Andrews, P., 1995. Experiments in taphonomy. Journal of Archaeological Science 22, David, F., 1997. Les ours du Châtelperronien de la Grotte du Renne a Arcy-sur-Cure 147e152. (Yonne). In: Tillet, T.H., Binford, L.R. (Eds.), L’Ours et l’Homme. Actes du Colloque Andrews, P., Fernández-Jalvo, Y., 1997. Surface modifications of the Sima de los d’Auberives-en-Royans, vol. 100. Eraul, Liège, pp. 185e192. Huesos fossil humans. Journal of Human Evolution 33 (2/3), 191e216. Domínguez-Rodrigo, M., 1994. Dinámica trófica, estrategias de consumo y alter- Arribas, A., Díez, J.C., Jordá, F.J., 1997. Primeras ocupaciones en los depósitos Pleis- aciones óseas en la sabana africana: resumen de un proyecto de investigación tocenos de la Cueva de los Torrejones (Sistema Central Español, Tamajón, etoarqueológico (1991e1993). Trabajos de Prehistoria 51 (1), 15e37. Guadalajara): litoestratigrafía y actividad biológica. Cuaternario y Geo- Domínguez-Rodrigo, M., 1999. Flesh availability and bone modification in carcasses morfología 11 (1e2), 55e66. consumed by lions. Palaeogeography, Palaeoclimatology, Palaeoecology 149, Auguste, P., 1991. Chasse et charognage au Paléolithique Moyen: l’apport du gise- 373e388. ment de Biache-Saint-Vaast (Pas-de-Calais). Bulletin de la Société Préhistorique Domínguez-Rodrigo, M., 2001. A study of carnivore competition in riparian and Française 80 (3), 68e69. open habitats of modern savannas and its implications for hominid behavioral Auguste, P., 1995. Chasse et charognage au Paléolithique Moyen: l’apport du gise- modelling. Journal of Human Evolution 40, 77e98. ment de Biache-Saint-Vaast (Pas-de-Calais). Bulletin de la Société Préhistorique Domínguez-Rodrigo, M., Rayne Pickering, T., 2003. Early hominid hunting and Française 92 (2), 155e167. scavenging: a zooarcheological review. Evolutionary Anthropology 12, 275e282. Azorit, C., Analla, M., Carrasco, R.J., Calvo, A., Múñoz Cobo, J., 2002. Teeth eruption Domínguez-Rodrigo, M., Piqueras, A., 2003. The use of tooth pits to identify pattern in red deer (Cervus elaphus hispanicus) in southern Spain. Anales de carnivore taxa in tooth-marked archaeofaunas and their relevance to recon- Biología 24, 107e114. struct hominid carcass processing behaviours. Journal of Archaeological Science Barone, R., 1976. Anatomie Comparée des Mammifères Domestiques. Vigot, París. 30, 1385e1391. Berger, G.W., Pérez-González, A., Carbonell, E., Arsuaga, J.L., Bermúdez de Castro, J.M., Domínguez-Rodrigo, M., Barba, R., 2006. New estimates of tooth mark and Ku, T.-L., 2008. Luminescence chronology of cave sediments at the Atapuerca percussion mark frequencies at the FLK Zinj site: the carnivore-hominid- paleoanthropological site, Spain. Journal of Human Evolution 55, 300e311. carnivore hypothesis falsified. Journal of Human Evolution 50, 170e194. Binford, L.R., 1981. Bones: Ancient Men and Modern Myths. Academic Press, New Emerson, A.M., 1993. The role of body part utility in small-scale hunting under two York, 320 pp. strategies of carcass recovery. In: Hudson, J. (Ed.), From Bones to Behavior: Binford, L.R., Mills, M.G.L., Stone, N.M., 1988. Hyena scavenging behavior and its Ethnoarchaeological and Experimental Contributions to the Interpretation of implications for interpretation of faunal assemblages from FLK22 (the Zinj Faunal Remains. University at Carbondale, Center for Archaeological Investi- floor) at Olduvai Gorge. Journal of Anthropological Archaeology 7, 99e135. gation, Southern Illinois, pp. 138e155. Occasional Paper, 21. Blasco, R., Rosell, J., 2009. Who was the first? An experimental application of Falguères, C., Bahain, J.J., Yokoyama, Y., Arsuaga, J.L., Bermúdez de Castro, J.M., carnivore and hominid overlapping marks at the Pleistocene archaeological Carbonell, E., Bischoff, J., Dolo, J.M.,1999. Earliest humans in Europe: the age of TD6 sites. Comptes Rendus Palevol 8, 579e592. Gran Dolina, Atapuerca, Spain. Journal of Human Evolution 37 (3/4), 343e352. Blumenschine, R.J., 1985. Early hominid scavenging opportunities: insights from the Fosse, P., 1994. Taphonomie paléolithique: les grands mammifères de Soleilhac ecology of carcass availability in Serengeti and Ngorongoro Crater, Tanzania. (Haute-Loire) et de Lunel-Viel 1 (Hérault). PhD Dissertation, Université de PhD Disertation. Berkeley, University of California. Provence, Aix-Marseille. Blumenschine, R.J., 1986a. Carcass consumption sequence and the archaeological Fosse, P., 1996. La Grotte n1 de Lunel-Viel (Hérault, France): repaire d'hyenes du distinction of scavenging and hunting. Journal of Human Evolution 15, Pleistocene Moyen. Paléo 8, 47e81. 639e659. Gaudzinski, S., Roebroeks, W., 2000. Adults only. Reindeer hunting at the Middle Blumenschine, R.J., 1986b. Early hominid scavenging opportunities: implications of Palaeolithic site Salzgitter-Lebenstedt, Northern Germany. Journal of Human carcass availability in the Serengeti and Ngorongoro Ecosystems. BAR Interna- Evolution 38, 497e521. tional Series 283, 163 pp. Giusberti, G., Peretto, C., 1991. Évidences de la fracturation intentionnelle d’osse- Blumenschine, R.J., 1988a. An experimental model of the timing of hominid and ments animaux avec moelle dans le gisement de "La Pineta" de Isernia (Molise), carnivore influence on archaeological bone assemblages. Journal of Archaeo- Italie. L’Anthropologie 95 (4), 765e778. logical Science 15, 483e502. Haynes, G., 1980. Evidence of carnivore gnawing on Pleistocene and recent Blumenschine, R.J., 1988b. Reinstating the early hominid scavenging niche: a reply mammalian bones. Paleobiology 6, 341e351. to Potts. Current Anthropology 29, 483e486. Haynes, G., 1983a. Frequencies of spiral and green-bone fractures on ungulate limb Blumenschine, R.J., 1995. Percussion marks, tooth marks, and experimental deter- bones in modern surface assemblages. American Antiquity 48, 102e114. minations of the timing of hominid an carnivore access to long bones at FLK Haynes, G., 1983b. A guide for differentiating mammalian carnivore taxa respon- Zinjanthropus, Olduvai Gorge, Tanzania. Journal of Human Evolution 29, 21e51. sible for gnaw damage to herbivore limb bones. Paleobiology 9, 164e172. Blumenschine, R.J., Selvaggio, M., 1988. Percussion marks on bone surfaces as a new Hill, A., 1984. Skeletal modification by modern spotted hyaenas. Abstracts. First diagnostic of hominid behavior. Nature 333, 763e765. International Conference on bone modification. Carson City, Nevada. pp.17e18. Bökonyi, S., 1972. Zoological Evidence for Seasonal or Permanent Occupation of Klein, R.G., 1975. Paleoanthropological implications of the nonarcheological bone Prehistoric Settlements, vol. 4. Warner Modular Publication. Reprint1e6. assemblage from Swartklip 1, South-Western Cape Province, South Africa. Brain, C.K., 1969. The contribution of Namib desert Hottentots to an understanding Quaternary Research 5, 275e288. of australopithecine bone accumulations. Scientific Paper of the Namib Desert Klein, R.G., 1989. Why does skeletal part representation differ between smaller and Research Station 39, 13e22. larger bovids at Klasies River Mouth and other archaeological sites? Journal of Brain, C.K., 1981. The Hunters or the Hunted? An Introduction to African Cave Archaeological Science 16, 243e256. Taphonomy. University of Chicago Press, Chicago, 365 pp. Lutman, F.C., 1993. Rhodesian Ridgebacks. TFH Publications, Inc., US. Bromage, T.G., Boyde, A., 1984. Microscopic criteria for the determination of Lyman, R.L., 1994. Vertebrate Taphonomy. Cambridge University Press, Cambridge, directionality cutmarks on bone. American Journal of Physical Anthropology 65, 550 pp. 359e366. Marean, C.W., Spencer, L.M., Blumenschine, R.J., Capaldo, S.D., 1992. Captive hyena Bunn, H.T., 1981. Archaeological evidence for meat-eating by PlioePleistocene bone choice and destruction, the schlepp effect, and Olduvai Gorge archae- hominids from Koobi Fora and Olduvai Gorge. Nature 291, 574e577. ofaunas. Journal of Archaeological Science 18, 101e121. Bunn, H.T., 1986. Patterns of skeletal representation and hominids subsistence Marean, C.W., Kim, S.Y., 1998. Mousterian large-mammal remains from Kobeh Cave. activities at Olduvai Gorge, Tanzania, and Koobi Fora, Kenya. Journal of Human Behavioral implications for Neanderthals and early modern humans. Current Evolution 15, 673e690. Anthropology 39, 79e113. Bunn, H.T., Bartram, L.E., Kroll, E.M., 1988. Variability in bone assemblage formation Mariezkurrena, K., Altuna, J., 1983. Contribución al conocimiento del desarrollo de la from Hadza hunting, scavenging, and carcass processing. Journal of Anthropo- dentición y el esqueleto postcraneal de Cervus elaphus. Munibe 35, 149e202. logical Archaeology 7, 412e457. Marshall, L.G., 1989. Bone modification and the laws of burial. In: Bonnichsen, R., Bunn, H.T., Harris, J.W.K., Isaac, G.L., Kaufulu, Z., Kroll, E.M., Schick, K., Toth, N., Sorg, M.H. (Eds.), Bone Modification. Center for the Study of the First Americans, Behrensmeyer, A.K., 1980. FxJj50: an early Pleistocene site in northern Kenya. Orono, Maine, pp. 7e26. World Archaeology 12, 109e136. Martínez Valle, R., 1996. Fauna del Pleistoceno Superior en el País valenciano. Capaldo, S.D., 1997. Experimental determinations of carcass processing by Plioe- Aspectos económicos, huellas de manipulación y valoración paleoambiental. Pleistocene hominids and carnivores at FLK 22 (Zinjanthropus), Olduvai Gorge, Ph.D Thesis. Universitat de València, Valencia, Spain. Tanzania. Journal of Human Evolution 33, 555e597. Menéndez, L., 2009. La Transición del Modo 2 al Modo 3 vista a través de la Capaldo, S.D., Blumenschine, R.J., 1994. A quantitative diagnosis of notches made by Industria Lítica de Gran Dolina TD10 (Atapuerca, Burgos) y Orgnac 3 (Ardèche, hammerstones percussion and carnivore gnawing on bovid long bones. Francia). Desarrollo técnológico y posibles implicaciones ocupacioneales de los American Antiquity 59 (4), 724e748. conjuntos. PhD Disertation. Rovira i Virgili University, Tarragona. Chauvière, F.-X., Castel, J.-C., 2004. Le statut du renard à Combe-Saunière (Sarliac- Michel, P., 2005. Un repaire würmien d’hyénes des cavernes: La Grotte d’Unikoté sur-l’Isle, Dordogne) et dans le Solutréen de l’Est d’Aquitain. In: Brugal, J.-P., (Iholdy, Pyrénées-Atlantiques, France). Museo de Altamira. Monografías 20, Desse, J. (Eds.), Petits Animaux et Sociétés Humaines. Du Complement 131e150. Alimentaire aux Ressources Utilitaires. XXIV Rencontres Internationales O’Connell, J.F., Hawkes, K., Blurton Jones, N., 1988. Hadza scavenging: implication d’Archéologie et d’Histoire d’Antibes. Editions APDCA, Antibes, pp. 389e402. for PlioePleistocene hominid subsistence. Current Anthropology 29, 356e363. Clot, A., 1987. La Grotte de la Carrière, repaire de Hyènes. In: Clot, A. (Ed.), La Grotte Oliver, J.S., 1993. Carcass processing by the Hadza: bone breakage from butchery to de Gerde (Hautes Pyrénées), Site Préhistorique et Paleontologique. Société consumtion. In: Hudson, J. (Ed.), From Bones to Behavior. Ethnoarchaelogical Ramond, Bagnères de Bigorre, pp. 191e207. and Experimental Contributions to the Interpretation of Faunal Remains. Cruz-Uribe, K., 1991. Distinguishing hyena from hominid bone accumulations. University at Carbondale, Center for Archaeological Investigations, Southern Journal of Field Archaeology 18, 467e486. Illinois, pp. 200e227. Occasional Paper, 21. 2060 R. Blasco et al. / Journal of Archaeological Science 37 (2010) 2051e2060

Parés, J.M., Pérez-González, A., 1999. Magnetochronology and stratigraphy at Gran Selvaggio, M.M., Wilder, J., 2001. Identifying the involvement of multiple carnivore Dolina section, Atapuerca (Burgos, Spain). Journal of Human Evolution 37 (3/4), taxa with archaeological bone assemblages. Journal of Archaeological Science 325e342. 28, 465e470. Pérez Ripoll, M., Morales Pérez, J.V., Sanchis Serra, A., Aura Tortosa, E., Sarrión Shipman, P., 1983. Early hominid lifestyle: hunting and gathering or foraging and Montañana, I., 2010. Presence of the genus Cuon in upper Pleistocene and initial scavenging? In: Clutton-Brock, J., Grigson, C. (Eds.), Animals and archaeology. Holocene sites of the Iberian Peninsula: new remains identified in archaeo- vol. 1. Hunters and their prey, vol. 163 British Archaeological Reports Interna- logical contexts of the Mediterranean region. Journal of Archaeological Science tional Series, Oxford, pp. 31e49. 37, 437e450. Shipman, P., Rose, J., 1983. Early hominid hunting, butchering and carcass-pro- Pickering, T.R., Egeland, C.P., 2006. Experimental patterns of hammerstone cessing behaviors: approaches to the fossil record. Journal of Anthropological percussion damage on bones: implications for inferences of carcass processing Archaeology 2, 57e98. by humans. Journal of Archaeological Science 33, 459e469. Shipman, P., Fisher, D.C., Rose, J., 1984. Mastodon butchery: microscopic evidence of Potts, R., Shipman, P., 1981. Cutmarks made by stone tools on bones from Olduvai carcass processing and bone tool use. Paleobiology 10 (3), 358e365. Gorge, Tanzania. Nature 291, 577e580. Silver, I.A., 1969. La determinación de la edad en los animales domésticos. In: Reitz, E.J., Wing, E.S., 1999. Zooarchaeology. Cambridge University Press, New York, Brothwell, D., Higg, E. (Eds.), Ciencia en Arqueología, s, Compiladores. Ciencia en 455 pp. Arqueología, Fondo de Cultura Económica, México D.F, pp. 229e239, pp. Riglet, P.-H., 1977. Contribution à l’étude de l’âge du Cerf élaphe (Cervus elaphus L.). 289e309. PhD Thesis, Ecole. Nationale Vétérinaire. Alfort, France. Stiner, M.C., 1994. Honor among thieves: a zooarchaeological study of Neanderthal Rodríguez Álvarez, X.P., 2004. Technical Systems of Lithic Production in the Lower ecology. Princeton University Press, Princeton, 447 pp. and Middle Pleistocene of the Iberian Peninsula. Technological Variability Sutcliffe, A.J., 1970. Spotted hyena: crusher, gnawer, digester and collector of bones. Between North-Eastern Sites and Sierra de Atapuerca Sites. BAR International Nature 227, 1110e1113. Series, Oxford. Tillet, T., 2002. Les grottes à ours et occupations néandertaliennes dans l’arc alpin et Rosell, J., 2001. Patrons d’Aprofitament de les Biomasses Animals durant el Pleistocè jurassien. In: Tillet, T., Binford, L.R. (Eds.), L’Ours et l’Homme. Actes du Colloque Inferior i Mig (Sierra de Atapuerca, Burgos) i Superior (Abric Romaní, Barcelona). d’Auberives-en-Royans, 1997, vol. 100. Eraul, Liège, pp. 167e184. PhD Thesis. Història i Geografia. Universitat Rovira i Virgili, Tarragona, Spain. Villa, P., Mahieu, E., 1991. Breakage patterns of human long bones. Journal of Human Rosell, J., Blasco, R., 2008. La presencia de carnívoros en conjuntos antrópicos del Evolution 21, 27e48. Pleistoceno Medio: el caso del nivel TD10-Sup de Gran Dolina (Sierra de Villa, P., Soressi, M., 2000. Stone tools in carnivore sites: the case of Bois Roche. Atapuerca, Burgos) y del nivel XII de la Cova del Bolomor (La Valldigna, Journal of Anthropological Research 56 (2), 187e215. Valencia). In: Díez, J.C. (Ed.), Zooarqueología Hoy. Encuentros Hispano-Argen- Villa, P., Castel, J.-C.H., Beauval, C., Bourdillat, V., Golberg, P., 2004. Human and tinos. Burgos University, Burgos, pp. 43e58. carnivore sites in the European Middle and Upper Paleolithic: similarities and Rosell, J., Blasco, R., 2009. Home sharing: carnivores in anthropogenic assemblages differences in bone modification and fragmentation. Revue de Paleobiologie 23 of the Middle Pleistocene. Journal of Taphonomy 7 (4), 305e324. (2), 705e730. Saitoti, T.O., 1988. The World of a Maasai Warrior. University of California Press, Yravedra, J., 2003e04. Interacción de humanos y carnívoros en el Pleistoceno Berkeley and Los Angeles, California, 144 p. Superior de la Península Ibérica. Novedosas interpretaciones en la Cueva de Saitori, T.O., Beckwith, C., 1980. Maasai. ELM Tree Books, London, 276 pp. Amalda. Espacio, Tiempo y Forma, Serie 1. Prehistoria y Arqueología 16e17, Selvaggio, M.M., 1994. Carnivore tooth marks and stone tool butchery marks on 79e98. scavenged bones: archaeological implications. Journal of Human Evolution 27, Yravedra, J., 2005. Aprovechamiento cárnico del lince (Lynx pardina) durante el 215e228. Pleistoceno superior en el interior de la Península Ibérica. Munibe 57, 303e311.