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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 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)

a r t i c l e i n f o a b s t r a c t

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
* Corresponding author. Tel.: þ34 977 257 882; fax: þ34 977559597.
carnivore to transport the skull (Cruz-Uribe, 1991). Furthermore,
E-mail address: rblasco@iphes.cat (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

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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).

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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
2001 excavations. Anatomical, taxonomic and modification 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
of Identified 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
formula was used: %survivali ¼ 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
The breaks on faunal remains were analysed and classified 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
Olympus SZ11 stereoscopic (magnification 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 3 2 1 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) were observed. Panthera leo 17 15 1 1
The analysis of cutmarks took into account the number of striations, Lynx sp. 1 1 1 1
Hystrix sp. 2 1 1 1
location on the anatomical element, distribution over the surface
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 1 1 1 1
Cervidae 121 24 2 1 1
lysed and the diagnostic elements of anthropogenic breakage were Megaloceros giganteus 1 1 1 1
documented on faunal remains from the TD10-1 sample: percussion Dama dama clactoniana 2 2 1 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
Hemitragus bonali 5 5 1 1
flakes (Capaldo and Blumenschine, 1994; Pickering and Egeland,
Capreolus capreolus 3 3 2 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

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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.

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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
Scapula 2
(2001), who studied the 1998e1999 excavation seasons from
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
The sample from TD10-1 (excavation seasons 2000e2001) 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
Calcaneus 1 1 2 50
(2.59%) belong to carnivores: U. arctos, C. lupus, V. vulpes, P. leo fossilis
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
a
related to lions (Fig. 2a, b). Minimal number of expected elements.

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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
Location Diaphysis Distal metaphysis The purpose of the lion capture is difficult 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
Measures (mm) 4.1e2.7 1.9e1.6
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

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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%)

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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) or the use of the
(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 1 1 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 anthropo-
genic 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
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