Steppe bison hunting in the Gravettian of Buda (lower Bistriţa Valley, eastern Romania)

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Original article

Steppe bison hunting in the Gravettian of Buda (lower Bistrit¸a Valley, eastern Romania) Chasse aux bisons des steppes dans le Gravettian de Buda (la basse vallée de la Bistrit¸a, Roumanie orientale) Valentin Dumitras¸cu a, S¸tefan Vasile b,* a b

Romanian Academy, ‘‘Vasile Pârvan’’ Institute of Archaeology, 11 Henri Coanda˘ Street, 010667 Bucharest, Romania University of Bucharest, Faculty of Geology and Geophysics, Department of Geology, Laboratory of Paleontology, 1 Nicolae Ba˘lcescu Avenue, 010041 Bucharest, Romania

Abstract The rich accumulation of bovid remains found at Buda (Baca˘u County, eastern Romania) is unique among the Upper Palaeolithic sites in the region. The morphological analysis of postcranial remains, which dominate the assemblage by far, shows they belong to Bison priscus, the steppe bison. The body parts representation is biased towards distal limbs, which also display typical butchery cut marks (resulted from disarticulation or skinning) and bone breakage, suggesting the presence of secondary butchery site where limbs were disarticulated and broken for marrow extraction. The assemblage is dominated by adults, mostly females, and includes few sub-adult individuals and no young juveniles. The large number of bison individuals indicates the found remains are the result of a mass killing event representing at least one hunting episode. Considering the age and sex structure of the population, we estimate that the event can be placed during autumn. # 2018 Elsevier Masson SAS. All rights reserved. Keywords: Butchering site; Seasonal hunting; Steppe bison; Gravettian; Eastern Romania

* Corresponding author. E-mail address: [email protected] (S¸. Vasile). https://doi.org/10.1016/j.anthro.2018.03.004 0003-5521/# 2018 Elsevier Masson SAS. All rights reserved.

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Re´sume´ L’accumulation riche de restes de bovidés trouvée à Buda (Comté de Baca˘u, Roumanie orientale) est unique parmi les sites du Paléolithique supérieur de la région. L’analyse morphologique des restes postcrâniens, qui domine, de loin, l’assemblage, montre qu’ils appartiennent à Bison priscus, le bison des steppes. La représentation des parties du corps est basée sur les membres distales, qui affichent aussi des marques de coupe typiques de l’abattage (résultent de pour la désarticulation ou de le dépouillage) et du cassage de l’os, suggérant qu’on est en présence d’un site d’abattage secondaire où les membres ont étés désarticulés et cassés pour l’extraction de la moelle. L’assemblage est dominé par des individus adultes, la plupart femelles, et comprend quelques individus d’âge sous-adulte, et pas du tout de juvéniles. Le grand nombre d’individus indique que les restes trouvés ont résulté d’un événement de tuerie en masse représentant au moins un épisode de chasse. Compte tenu de l’âge et de la distribution sexuelle de la population, nous estimons que l’événement peut être situé pendant l’automne. # 2018 Elsevier Masson SAS. Tous droits réservés. Mots clés : Site d’abattage ; Chasse saisonnière ; Bisons des steppes ; Gravettien ; Roumanie orientale

1. Introduction One of the largest rivers in Moldova (eastern Romania), Bistrit¸a crosses the Eastern Carpathians from north-west to south-east, facilitating the access across the mountain range. Evidence of Palaeolithic human habitation is documented for both the upper (mountainous) and the lower (hilly) sectors of the valley, including well-documented sites such as BistricioaraLuta˘rie, Bicaz, Dârt¸u, Poiana Cires¸ului, Buda and Lespezi (Nicola˘escu-Plops¸or et al., 1961a, 1961b; Ca˘pitanu et al., 1962; Bitiri and Ca˘pitanu, 1972; Bolomey, 1989; Bitiri-Ciortescu et al., 1989; Pa˘unescu, 1998, 1999; Cârciumaru et al., 2008; Nit¸a˘-Ba˘la˘s¸escu, 2008). Most sites are located upstream, in the Ceahla˘u Basin, but the largest amount and the best preservation of animal bones is recorded from the group of three sites situated downstream on the river valley: Buda, Lespezi, and Poiana Cires¸ului. The latter two were interpreted as seasonal residential sites, with rich lithic and faunal assemblages dominated by reindeer (Poiana Cires,ului) or reindeer and horse (Lespezi). For both these sites the species diversity is quite large, the skeletal representation of the animals is complete, bone and antler industry is present and also habitation structures and consistent fireplaces were identified. (Bolomey, 1989; Pa˘unescu, 1998; Beldiman, 1999; Dumitras¸cu, 2008). The other Palaeolithic site, Buda, was discovered in 1958 (Ca˘pitanu et al., 1962; Nicola˘escu-Plops¸or et al., 1961a) on a hilltop northwest of Buda village, Baca˘u County, a site known locally as ‘‘Dealul Viilor’’, overseeing the wide lower Bistrit¸a Valley. Here, large mammal fossil remains occur alongside lithic elements in a 40-cm thick layer, suggesting they belonged to the same cultural event (Nicola˘escu-Plops¸or et al., 1961a; Ca˘pitanu et al., 1962; Tuffreau et al., 2018). The lithic elements show a typology characterizing the Gravettian, the absolute chronology also placing the cultural layer during this cultural unit – radiocarbon dating of charcoal and bone fragments estimated the age at 23.810  190 years BP, (Pa˘unescu, 1998, but also see Tuffreau et al., 2018). Bone and antler industry is absent, no consistent habitation structures were found, fire traces are scarce and dispersed, and the body parts representation of the animals is overwhelmed by distal limb elements. The faunal remains – more than 1000 – unearthed during the first excavation (1958–1962) were found as bone agglomerations mainly consisting in long bone extremities, complete small bones and a few jaw fragments attributed to

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the aurochs (Bos primigenius) and reindeer (Rangifer sp.) (Bolomey, 1961). Another 1020 remains resulted from the following excavations, of which 900 were determinable (Necrasov and Bulai-S¸tirbu, 1972). Out of these, almost 90% were assigned to large bovids, most probably bison, followed by the reindeer (about 10%), and less than 1% to horse and red deer. Bolomey (1966) mentions that some postcranial bones from the Palaeolithic sites along Bistrit¸a Valley cannot be confidently assigned to the aurochs, since they present features comparable to those of the bison (the steppe bison, Bison priscus). In consequence, subsequent syntheses regarding Palaeolithic sites from this area usually refer to the large bovid remains as Bos/Bison (e.g., Pa˘unescu, 1998). The apparent concentration of bones on some areas of the site, and also the prevalence of metapodials and phalanges was considered to be the result of a magical ritual, performed in order to summon the animals. However, long bone breaking for marrow removal is recognized in early works (Bolomey, 1961; Nicola˘escu-Plops¸or et al., 1961a; Ca˘pitanu et al., 1962; Ca˘pitanu, 1967). 2. Material and methods The megafaunal remains analyzed in this study were recovered during the initial excavation of the site (Nicola˘escu-Plops¸or et al., 1961a; Ca˘pitanu et al., 1962), but also during recent excavations (2014–2015) carried out by the ‘‘Vasile Pârvan’’ Institute of Archaeology in collaboration with Lille 1 University – Science and Technology, within the frame of the Archaeological Mission ‘‘The Palaeolithic of Romania’’. The specimens from the initial excavation are housed at the National History Museum of Romania, Bucharest (MNIR) and at the ‘‘Iulian Antonescu’’ Museum Complex, Baca˘u, whereas the specimens from recent excavations (more than 600 specimens) are kept at the ‘‘Vasile Pârvan’’ Institute of Archaeology, Bucharest (IAB). A total of 1229 faunal remains were studied, both morphologically and morphometrically. Only the fragments large enough to preserve some morphological element that allowed deduction of their anatomical position (1044 specimens) were analyzed in this paper, the rest representing long bone shaft splinters. Taxonomical assessment of the fossil remains is based on the works of Olsen (1960), Gee (1993), and Sala et al. (2010). Anatomical attribution of complete or fragmentary bones was made by direct comparison to extant ox bones from the IAB reference collection, and was also based on Pales and Lambert (1971). Terms describing various aspects of bone morphology follow Olsen (1960). Morphometric parameters employed in estimating the gender ratio were measured according to Driesch (1976), using a 0.01 mm precision digital calliper. The outline of a Bison bonasus drawn by Michel Coutoureau and made available on http:// www.ArcheoZoo.org was used to represent the skeletal parts of the assemblage and the areas where bone marks are present. 3. Taxonomic assessment Late Pleistocene megafaunas of Europe are characterized by the presence of two large bovids: the steppe bison (Bison priscus) and the aurochs (Bos primigenius) (e.g. Kahlke, 1999; Vasiliev, 2008; Petronio et al., 2011). The two taxa are also quite common during the late Pleistocene of Romania, where they occur in both natural (Apostol, 1967, 1976; Radulesco and Hermann, 1971; Apostol and Cacoveanu, 1980; Codrea and Ursachi, 2010) and archaeological (e.g., Pa˘unescu, 1998, 1999) contexts.

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Discerning between the two late Pleistocene large bovid taxa proved to be difficult for the bovid remains excavated from Buda, since cranial remains are extremely rare, and, when present, they are much to fragmentary to present relevant features for a precise taxonomical assessment. Bolomey (1961) first assigned the bovid postcranial elements from Buda to Bos primigenius, without offering many arguments for this conclusion. Later on, she recognizes that not all the bovid material from several sites on the lower Bistrit¸a Valley can be positively assigned to Bos primigenius, some specimens showing morphological features indicative for Bison (Bolomey, 1966). In order to address this matter, all postcranial elements that were complete enough to preserve relevant features underwent a thorough inspection of their morphology. The results were compared to the observations made by Olsen (1960), who compared the morphological differences between extant Bos and Bison postcranial elements. In doing so, we considered that the morphological differences usually present at the level of bone epiphyses between Bos taurus and Bison bison are relevant enough to be used as proxy for inter-genus differentiation, and, therefore, also applicable to discerning between fossil species of the same genera: Bos primigenius and Bison priscus, conclusion also reached by other authors (e.g. Gee, 1993; Sala et al., 2010). Additionally, the fossil material from Buda was compared to extant Bos taurus material, from the reference collection of the IAB. The morphological features used for taxonomical assessment are mentioned here only for some of the most abundant elements that were complete enough to allow the evaluation of the characters (Table 1): distal tibiae, distal humeri, astragali, and calcanei (Fig. 1). In distal view, the tibiae of Bos and Bison differ mainly in the morphology of the articulation surface for the malleolus: in Bos the anterior articulation surface for the malleolus is mediolaterally compressed and antero-posteriorly elongated, lower, and confluent with the posterior one (Fig. 1–1); in Bison the anterior articulation surface for the malleolus is rather circular, more raised (i.e. distally extended) and clearly separated from the posterior one by a deep notch; (Fig. 1–2) (Gee, 1993; Sala et al., 2010). In anterior view, the astragali of Bos and Bison differ in the position of the medial tubercle relative to the common tangent to the proximal margins of the distal trochlea: in Bos the tubercle always falls below (i.e. distal to) the line (Fig. 1–3), whereas in Bison the tubercle is located above (i.e. proximal to) this line, or, at most, is crossed by it (Fig. 1–4) (Olsen, 1960). In medial view, the relative position of the medial condyle and medial epicondyle of the distal humerus differs between Bos taurus and Bison: the medial epicondyle clearly projects downward (i.e. more distally) relative to the medial condyle in Bos taurus (Fig. 1–5), whereas in Bison the distal ends of the two elements are placed almost in the same plane, the distal end of medial epicondyle being found only slightly distally from the distal end of the medial condyle (Fig. 1–6) (Olsen, 1960). In medial view, the calcaneum of Bos taurus differs from that of Bison in the shape of the sustentaculum, which has two almost perpendicular edges, and, therefore, a more squarish aspect in the former (Fig. 1–7), but appears as rounded in the latter (Fig. 1–8) (Olsen, 1960). None of the elements complete enough to be considered relevant in the morphological analysis showed morphological features that would indicate assignment to Bos (in general) or, by extension, to Bos primigenius. Therefore, we consider that the entire sample of large bovid postcranial skeletal elements, fragmentary or complete, can be confidently assigned to the genus Bison, and, by extension, to Bison priscus, the only bison species commonly mentioned from the area during the late Pleistocene (but see Soubrier et al., 2016 for a different opinion, suggesting that another bison species, deemed as the direct ancestor of the European wisent, Bison bonasus, was also present at the time).

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Table 1 Anatomical distribution of the skeletal elements used in this study. The number in parentheses shows how many from the total number of elements had unfused epiphyses. Distribution anatomique des éléments squelettiques utilisés dans cette étude. Le nombre entre parenthèses montre combien d’éléments ont eu des épiphyses non fusionnées par rapport au nombre total. Anatomical element

Right

Left

Indeterminate

Total

Cranium Isolated teeth Mandibula Rib Scapula Humerus – proximal Humerus – distal Radius – proximal Radius – distal Ulna – proximal Scaphoid Semilunar Pyramidal Pisiform Unciform Capitato–trapezoid Metacarpal – proximal Metacarpal – distal Coxal Femur – proximal Femur – distal Femur – shaft Patella Tibia – proximal Tibia – distal Malleolar Astragalus Calcaneus Cuneiform Sesamoid Metatarsal – proximal Metarasal – distal Metapodial – distal Phalanx I – anterior Phalanx I – posterior Phalanx I – indeterminate Phalanx II – anterior Phalanx II – posterior Phalanx II – indeterminate Phalanx III

– – 5 – – 1 21 (2) 35 12 (4) – 9 10 6 1 7 8 13 3 – 9 5 (2) – 3 16 (1) 41 (5) 12 25 16 (3) 3 – 15 8 – – – – – – – –

– – 2 – – 5 4 (1) 21 23 (6) – 12 9 5 2 13 9 8 7 (1) – 10 8 – 6 7 (2) 28 (5) 13 38 12 (1) 2 – 18 15 – – – – – – – –

2 27 7 11 7 6 (3) 8 (3) – 5 (5) 3 – – – – – – – 9 5 2 (2) 3 12 – 6 5 (1) 1 – 1 – 20 – – 72 (41) 42 55 13 (12) 9 8 130 (1) 21

2 27 7 11 7 12 (3) 33 56 40 (15) 3 21 19 11 3 20 17 21 19 (1) 5 21 (2) 16 (2) 12 9 29 74 (11) 26 63 29 (4) 5 20 33 23 72 (41) 42 55 13 (12) 9 8 130 (1) 21 1044 (98)

4. Skeletal part representation The representation of various skeletal elements in a natural or archeological site is related to the processes that affected the animal’s body after its death and prior to its burial in sediment, or, in other words, by its taphonomical history (Efremov, 1940; Lyman, 1987, 1994). A complete

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Fig. 1. Morphological differences between some of the postcranial elements of the ox (Bos taurus) and of the steppe bison (Bison priscus) from Buda: distal tibiae of ox (1) and steppe bison (2), in distal views; astragali of ox (3) and steppe bison (4), in anterior views; distal humeri of ox (5) and steppe bison (6), in medial views; calcanei of ox (7) and steppe bison (8), in medial views. See text for details on the morphological elements indicated by lines and arrows. Images not to scale. Différences morphologiques entre quelques éléments postcrâniens du bœuf (Bos taurus) et du bison des steppes (Bison priscus) de Buda : tibias distales de bœuf (1) et de bison des steppes (2), en vues distales; astragales de bœuf (3) et de bison des steppes (4), en vues antérieures ; humérus distales de bœuf (5) et de bison des steppes (6), en vues médiales ; calcanéums de bœuf (7) et de bison des steppes (8), en vues médiales. Voir le texte pour les détails sur les éléments morphologiques indiqués par des lignes et des flèches. Les images ne sont pas à la même échelle.

skeleton found in articulation indicates the animal’s carcass was not disturbed after death, but such cases are rather rare. Weathering of fragile bones, hydrodynamic transport of skeletal remains during fluvial sedimentation, carcass or bone scavenging by carnivores or even other herbivores, trampling by animals, or, in the case of archaeological assemblages, anthropic activities, are all factors that contribute to preferential accumulation of various skeletal elements. The bone assemblage from Buda is overwhelmingly dominated by elements of the appendicular skeleton: out of 1044 elements that could be assigned to an anatomical element (Table 1), only 59 (5.65% of all bones) were identified as not belonging to limb bones (small pieces of skull, dental, mandibular, scapula, rib and coxal fragments). Therefore, it becomes obvious that some selective transport agent created the bone assemblage from Buda. Weathering can also contribute to bone fragmenting, leading to the preferential preservation of the most mineralized parts of the skeleton. Teeth, the most mineralized body parts, and, therefore, the most resistant to weathering are not present in high number in the bone assemblage from Buda – only 27 dental fragments (2.58% of the assemblage) were recovered, showing that weathering did not play an important part in postcranial bone fragmentation.

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The bones and stone tools from Buda were found in a layer of yellow-brownish silty sediment, of loessoid appearance. The sediment is well sorted, and it contains large pores filled with carbonate calcretes deposited in the vadose zone by water evaporation. A network of thin millimeter-wide shallow grooves is usually present at bone surface, at the level of the periosteum, and might represent root etchings (Lyman, 1994). Small-scale dendrites of manganese oxides are also quite common on bone surfaces. Bone fractures are sharp, showing they were not rolled by water. Although no detailed sedimentological study was yet performed to confirm it, the elements mentioned above support an aeolian sedimentation for the layer that yielded the faunal remains from Buda. A few gastropod shells were also found in the sediment. The preliminary taxonomical determination of the gastropods (personal observation, unpublished data) indicates they belonged to terrestrial taxa (Vertigo sp., Succinea sp., Vallonia sp.), which also supports a subaerial accumulation of the yellow-brownish silt. Since no evidence of fluvial sedimentation was found, hydraulic transport can be ruled out as factor of preferential accumulation of certain skeletal parts. Since neither carnivore bites nor chew marks were observed on any of the bones from Buda, scavenging of the carcasses is not supported either. A single bone presented gnawing marks, but their morphology is not consistent with carnivore activity (see below, section 5. Bone modifications). Of the limb bones, the best represented skeletal region, none of the long bones are complete. They are only present as fragmentary extremities (more or less complete epiphyses), or as diaphyseal fragments. The only elements that were also found at Buda as complete bones are the carpals, tarsals, phalanges, and kneecaps. Where possible, the limb bones or their extremities were also separated according to bilateral symmetry into left and right elements, in order to estimate the minimum number of individuals (MNI) present at the site. The most abundant elements are right distal tibias, which allow the estimation of MNI to 41. The steppe bison elements present in the bone assemblage from Buda are listed in Table 1 and illustrated in Fig. 2. Since only the lateral left side of a bison is viewable, the elements figured are not lateralized (e.g. all the left, right and laterally indeterminable elements are figured on the left side). Because numerous traces of anthropic activity were found on the limb elements (see below, section 5. Bone modifications), the high amount of limb bones from Buda is related to anthropic causes, and, more precisely, to butchering activities. In the case of large mammals, where transporting a whole carcass that weighs several hundred kilograms is practically impossible, butchering must take place at the kill site. According to the distance the parts of the carcass need to be transported across, some body parts that offer more food resources (meat, fat, marrow) are selected and transported, whereas other, bulkier and offering less food, are abandoned – a selective transport decision named the ‘‘schlepp effect’’ by Perkins and Daly (1968). Various strategies were observed in the hunting behaviour of present-day hunter-gatherer societies (e.g. Binford, 1978; Bunn et al., 1988; O’Connell et al., 1990; Lupo, 2001) and used to construct models by which to interpret the hunting and butchering behaviour of prehistoric hunters, based on information obtained from archaeological bone assemblages (e.g. White, 1952, 1953; Perkins & Daly, 1968; Metcalfe & Jones, 1988; Emerson, 1993). The general interpretation is that archaeological bone assemblages including remains of large mammals that were hunted (in opposition to domesticated) are dominated by elements of the axial skeleton (skull, vertebrae) most likely represent primary butchering sites near the kill site. On the other hand, when the assemblage is dominated by limb remains, it most likely represents a secondary butchering site, on route to, or at a temporary or more permanent camp. However, not all parts of the limbs contain meat–the distal limbs, including the ankles, metapodials and phalanges offer very little

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Fig. 2. Skeletal parts of steppe bison from Buda. Dotted line is used for elements found only as fragments, and solid line outline is used for elements that were also found complete. Parties squelettiques du bison des steppes de Buda. La ligne en pointillée est utilisée pour les éléments trouvés seulement en fragments, et la ligne continue pour des éléments qui ont été trouvés complets.

meat, yet they occur in some bone assemblages. The decision to transport such elements (along with other limb bones) seems to be based not solely on meat content, but also on the fat content of bone marrow, which can be extracted even from metapodials (Bunn et al., 1988; Emerson, 1993). Perkins and Daly (1968) suggest that distal limb elements are transported along with meat- and marrow-bearing long bones also because they offer a firm grip and make it easier for the entire limb to be carried to camp. Some elements chosen for transport might serve as ‘‘snacks’’ on the way from the kill site to camp (Bunn et al., 1988). As mentioned above, no long limb bone was found intact, and many of the epiphyseal fragments bear impact points (see below, section 5. Bone modifications). Charred epiphyseal fragments, knuckle bones and phalanges were also found, showing that some of the prey was cooked and consumed at the site. The dominance of limb elements in the bone assemblage from Buda fits the model of a secondary butchering site, where marrow extraction took place. 5. Bone modifications The stone tools and hearths found at Buda are not the only evidence of human activity at the site. Many bone fragments bear the marks of anthropic activity, the most frequent being traces of impact, cut marks, and the presence of charred bones or bone fragments. Next to the striking dominance of one large bovid taxon, another aspect noticed from the time of the first excavations was the absence of complete long bones – save for the smaller and compact carpal and tarsal bones and the phalanges, none of the limb bones were found intact, but

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as fragments, either consisting in epiphyseal regions or in small diaphyseal shards and splinters. Except for one case of articulated distal limb (all six phalanges, from proximal to distal) there are no other examples of elements in anatomical connection. The most common marks found on the bones from Buda are impact points, associated with spiral cracks. Often times, more impact points can be seen close to each other on the same bone, suggesting that bone breaking was not accidental, but a clear activity in carcass processing. As noted above (section 4. Skeletal part representation), meat content is not the only deciding factor in selecting what body parts are to be transported from the kill site, the decision being also influenced by bone marrow content (e.g. Binford, 1978; Metcalfe & Jones, 1988; Emerson, 1993). Although spiral cracks occur on fresh bone in general (Binford, 1978), and are not necessarily linked to marrow extraction, the occurrence of multiple impact points, aimed to increase the length of the cracks, is evidence supporting the latter activity (Kooyman, 2004). However, not all impact points leading to bone breaking are linked to marrow extraction. Separating the limbs from the torso can also be done by breaking the proximal ends of humeri and femora, but it is considered that this activity produces less impact points on bone surface than marrow extraction (Kooyman, 2004). Disarticulation of the humerus and femur by smashing their proximal epiphyses leaves the proximal ends connected to the thoracic or pelvic girdle, both abandoned at the primary butchering site. Since some proximal humeral and femoral ends are present (albeit in relatively lower number), alongside some coxal and scapular fragments, we conclude that joint dismemberment was made by smashing the latter elements, which left the femur and humerus intact after primary butchering and transportable as complete elements. Therefore, the breaking of femora and humeri must have taken place as a secondary butchering activity, for marrow extraction. Impact points are present on the broken limb bones from Buda, and they are associated with spiral fractures, suggesting that anthropic bone breaking occurred at the site, as part of the butchering process. All long limb bones present impact marks, observed more frequently on tibias and metapodials (Figs. 3 and 4). However, this observation might also be biased by the much higher abundance of latter elements in the assemblage (Table 1, Fig. 2). Impact points occur on both the anterior and posterior side, next to proximal or distal ends of tibias, ulnas and metapodials, with no preferential distribution (Fig. 4). Fine subparallel grooves were seen on several bones, and interpreted as cut marks that can be produced during primary or secondary butchering activities, in order to remove the hides, strip the meat off the bones, or cut tendons and ligaments during joint dismemberment. Disjointing cut marks were identified on several anatomical regions: the coxo-femoral, humero-radial, tibio-talar and metapodial-phalangeal articulations. Most cut marks were found on the medial and anterior sides of astragals (Fig. 3–6 and Fig. 3– 7), and were most probably made during joint dismemberment. Other cut marks, present on proximal metapodials, might also be the result of dismembering, whereas cut marks found on phalanges and proximal diaphyses of metapodials are probably the result of skinning. One proximal femur fragment also bears cut marks, possibly as result of meat stripping. Charred bones are also common at the site, including distal ends of humerus and metatarsal, proximal and distal ends of tibia and metacarpal, but also complete phalanges and knuckle bones (astragal and unciform) (Fig. 3–8,Fig. 3–9, Fig. 3–10). Ash patches, representing the remainders of hearths are also present at the site, and show that thermal processing must have taken place, either for meat or for in-bone cooking of marrow. Anthropic traces are not the only ones found on the bones–although no sign of carnivore activity was found, a single astragalus presented a series of closely packed grooves along its

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Fig. 3. Modifications on steppe bison bones from Buda: impact points on broken tibial distal ends, in posterior (1) and anterior (2–4) views; impact points and cut marks on broken proximal end of metacarpus (5), in posterior view; cut marks on complete astragali (6–7), in medial views; charred distal tibia (8), in posterior view, astragalus (9), in anterior view, and second phalanx (10), in lateral? view; rodent gnawing marks on anteromedial edge of astragalus (11). Images not to scale. Modifications sur les os de bison des steppes de Buda : points d’impact sur des extrémités distales de tibias cassés, en vues postérieure (1) et antérieure (2–4) ; points d’impact et marques de coupe sur l’extrémité proximale d’un métacarpe cassé (5), en vue postérieure ; marques de coupe sur des astragales complets (6–7) en vues médiales ; tibia distal brûlé (8), en vue postérieure, astragale brûlé (9), en vue antérieure, et seconde-phalange (10), en vue latérale? ; marques de rongeur sur la marge antéromédiale d’un astragale (11). Les images ne sont pas à la même échelle.

anteromedial margin (Fig. 3–10). The grooves are about 1 mm deep and about 5 mm wide, and correspond to the pattern left by the spatulated incisors of rodents. Rodent gnawing marks on bones are not uncommon, and were assigned to various taxa (e.g., Wood, 1952; Duthie and Skinner, 1986; Klippel and Synstelien, 2007). Observations on present taxa show that some rodents gnaw on bones for nutrients contained in their fat, whereas others do it for the minerals contained in the bone, after the grease had leached away (Klippel and Synstelien, 2007). Recent observations suggest that some rodents are not gnawing on bones for feeding purposes, but as a behavioural adaptation, in order to hone and trim their life-long continuous growing incisors (Kiibi, 2009). Although rodents of various sizes are known from Palaeolithic sites across the

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Fig. 4. Distribution of bone modifications observed in the steppe bison specimens from Buda. Distribution des modifications sur les os observées chez les spécimens de bison des steppes de Buda.

country (e.g. Paunesco et al., 1982), the gnawing marks are from Buda cannot be assigned to a certain taxon.

6. Age and sex representation Dental eruption sequence and tooth wear stages are the best indicators for the age of an animal. Very few teeth were found, so, to estimate the age structure of the population, the stages of epiphyseal union of the long bones and the relative dimensions of the skeletal elements were used. The remains of immature individuals (unfused epiphyses) are dimensionally comparable to the adult bones indicating a sub-adult degree of ontogenetic development (Bement, 1999). Considering the element used for estimating the MNI and sex distribution, the right distal tibia (see below), the amount of immature yet full body sized individuals is around 12%. Taking the moment of epiphyseal fusion of various bones into account, the only elements indicating the presence of yearlings (two individuals less than 1.3 years) are the distal humerus and second phalanx (Bement, 1999). No elements from very young animals (calves) were identified. In absence of cranial material, the best for sexing bison remains from natural or archaeological assemblages (Reher, 1970; Speth, 1983), some of the most useful postcranial remains that can supply relevant information in this respect are distal limb bones. Bison metapodials are relevant since they must support much more weight in the case of males than in females. Not only are the males larger overall, but their limbs (the anterior ones in particular) must also offer supplementary strength and stability during head-butting. Since some degree of overlap between younger males and larger females may exist, a multivariate analysis should be used for sexing bison metapodials (e.g., Lewis et al., 2005). However, complete metacarpals or metatarsal are not

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a common find in archaeological sites, since they are many times broken for marrow extraction (Emerson, 1993; Kooyman, 2004), or used as tools. Indeed, no complete metapodials were found at Buda, the most common elements recovered being their proximal and distal ends. In order to avoid counting the same individual twice, and thus artificially increasing the ratio of that gender in the sample, only lateralized elements (i.e. positively identified as belonging to the left or right side) must be used. Limb bones presenting incompletely fused epiphyses, therefore belonging to individuals that did not reach adulthood, were also eliminated from the sample used for gender assessment, since elements of subadult males and those of females would overlap in size. In consequence, the remaining number of adult measureable metapodial ends belonging to the same side was quite low, so a different method had to be used in order to address the sexual dimorphism for the bison assemblage of Buda. Since they are the most numerous bones in the sample, and they relate to the metatarsals via tarsals, therefore having to accommodate weight in a similar manner, and thus accordingly increasing or decreasing in size, distal ends of tibia were used to estimate gender ratio. More right measurable distal tibias with completely fused epiphyses were identified in the sample, and are here used to estimate gender ratio. Because no complete tibiae were found, a multivariate representation could not be used, so a bivariate analysis was used in this study, taking into account the depth (i.e. antero-posterior diameter) and breadth (i.e. mediolateral diameter) of the distal tibial ends (Fig. 5). The results obtained by plotting the measurements of tibial distal ends indicated that 66% of the samples represent female individuals, whereas only 33% represent males. Although in present populations the number of males is sometimes even higher than the one of females during the growing season, and might vary significantly between different groups (Prezanowski et al., 2012), an estimate of the sex ratio proposed by Krasin´ska and Krasin´ski (2013) for a larger, stable and self-sustaining population of European bison (Bison bonasus) is similar to the one obtained from the bone assemblage of Buda. A comparable ratio (1:2 ratio between male and female bison) was reported for mixed groups of Bison bison with a slight increase in the percentage of males during rut intervals (e.g. Fuller, 1960; Komers et al., 1992). The matter of estimating the yearly rates of ontogenetic development in Bison priscus is quite delicate, since the species is extinct for more than 12 000 years. The two extant bison species, Bison bison and Bison bonasus, can be used as proxies, although it is debatable which approaches their extinct relative best in behaviour. Both the European and the American bison breed in late

Fig. 5. Measurements of left distal tibiae used for sex assignment of the steppe bison from Buda. Circles represent cows and squares represent bulls. Mesures des tibias distales gauches employées pour l’attribution des bisons des steppes de Buda par sexe. Les cercles représentent des vaches et les carrés des taureaux.

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summer–early autumn (Haugen, 1974; Krasin´ska and Krasin´ski, 2013, Chapter 12. Reproduction), which results in most births happening during mid–late spring. According to these dates, a population that includes adults and sub-adult juveniles would correspond to mid– late autumn. However, these estimates should be used with caution, since the breeding season, and, therefore the birthing interval as well, might extend over a longer time interval (Walde, 2006). Also, since the annual life cycle of wildlife depends on climate, information about the succession of seasons and annual mean temperatures during the exact period in which the steppe bison from Buda lived must also be taken into account, as well as the pattern of their possible migrations–all these aspects remain, for now, unknown.

7. Hunting strategies and bison behaviour There are two main factors that can explain the missing calves from the archaeological bison population: 1. the taphonomical conditions did not allow the preservation of their fragile bones; 2. the animals were not killed during the calving period, the spring. For the new excavations, dry and wet sieving was used to retrieve small objects and microfauna. Even though in the old excavations the sieve was not employed, small bones and fragments were nonetheless recovered. The preservation state is sufficient to conserve small and fragile elements, so the absence of young individuals cannot be the consequence of differential preservation or the selection of bones during the excavation. As a comparison, the faunal assemblages from the neighbouring sites, Lespezi and Poiana Cires,ului, recovered from comparable loess deposits, contain elements from very young individuals (reindeer), including new born and foetal bones (Dumitras,cu, 2008). The animals identified in the archaeological assemblage represent a sample extracted from the living population. Is this sample an accurate reflection of the living population or it does represent a selection biased toward particular categories of animals? If there is a selection, was it induced by the hunters’s strategies, the bison’s behavior, and herd structure or by a combination of different variables? Selective hunting implies targeting the individuals that are best suited for the needs of humans in a specific time of the year. If we follow this hypothesis than we conclude that the hunters from Buda preferred adult females, so we can place the hunting event (or events) in the fall/early winter, when cows are in good shape, whereas bulls are in bad condition after the mating season, there are no young calves, and the individuals born in the spring are already grown to near-adult size (Speth, 2013 and references therein). If the hunt was not selective, then the archaeological assemblage must reflect the structure of the living bison population at the time of the kill. Again, the resulting age/sex structure of the archaeological population indicates the end of the growing season/beginning of the cold season. The animals might have been ambushed, or driven in a natural (e.g. cul-de-sac, precipice) or manmade trap. The bison herd behavior when attacked by a pack of predators is to form a running column with a specific age and sex structure: the adult females generally group in front, and the calves follow them, while some of the adult males remain in the back, to fend off predators. If the column is driven towards a trap or ambush, the animals from the front will mostly be killed, the ones from the rear having the chance to escape. Thus, the predominance of adult females and the under-representation or absence of adult males and calves in some archaeological assemblages may not necessarily indicate a fall/early winter kill event (Speth, 1997, 2013). At Buda there is a dominance of adult females, but it cannot be said that bulls are under-represented, since they

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occur in the bone assemblage in ratios close to those recorded for natural populations (33% of total). In addition, a number of adolescents are present and calves are the missing category. According to the age and sex structures of the steppe bison assemblage from Buda, similar to those occurring in natural aggregate populations during or shortly after mating season, it seems most probable that the accumulation is the result of an unselective mass kill taking place around autumn. 8. Conclusions The thorough morphological analysis of the large bovid assemblage from Buda shows that the main target of the Gravettian hunters from this site was the steppe bison (Bison priscus). We don’t know yet if the steppe bison from Buda was migratory or resident. In any of the two cases, the significant number of animals (over 41) and the sex structure of the assemblage place the kill in a period of aggregation, which occurs in the rut period, during autumn. The age structure of the assemblage, dominated by adults, including a few yearlings and devoid of calves, also indicates the hunting did not take place during spring or summer, the time of calving. The large number of animals suggests a communal mass kill, where little selection of individuals by sex or age is possible; therefore we incline to consider that the archaeological population reflects in a reasonable measure the living population from which it was extracted. The absence of axial skeleton elements, bulkier and containing less food resources, suggests they were left at the primary butchery site. The assemblage consists almost exclusively of limb transported from the kill site because of their higher meat and marrow content. Traces of impact and cut marks found on bones also indicate the context of a secondary butchering site, where marrow was retrieved from all the long bones, and where skins were also recovered. All that was left at the site were low utility elements (distal limbs), fragments and splinters from the broken long bones. This picture suggests a temporary camp were the secondary butchery of bison was conducted, following successful hunting, in preparation for the cold season. Acknowledgments The authors are grateful to Dr. Roxana Dobrescu (IAB, Bucharest) and Dr. Alain Tuffreau (Lille) for inviting them to take part in the excavations and for the information supplied on the cultural archaeological context of the Buda site. Dr. Valentin Radu (MNIR, Bucharest) is thanked for facilitating access to the specimens stored in the collections of the MNIR. Dr. Adrian Ba˘la˘s¸escu (MNIR, Bucharest) is thanked for the discussions regarding various aspects of archaeozoology and for providing us with helpful literature. References Apostol, A., 1967. Étude des bovidés quaternaires de la région de Bucarest. Travaux du Muséum national d’histoire naturelle « Grigore Antipa », 7. pp. 449–462. Apostol, A., 1976. L’étude morphométrique de mammifères fossiles quaternaires de la plaine Roumaine et leur distribution paléozoogéographique. Travaux du Muséum national d’histoire naturelle « Grigore Antipa », 17. pp. 341–375. Apostol, A., Cacoveanu, I., 1980. L’étude des restes fossiles quaternaires d’Eléphantidés, de Bovidés et de Cervidés conservés dans les musées des villes d’Alexandria et de Ros¸iorii de Vede (Département de Teleorman, Roumanie). Travaux du Muséum national d’histoire naturelle « Grigore Antipa », 22. pp. 587–607.

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