The article presents the results of a detailed zooarchaeological and taphonomic study of materials from the Bondi Cave in Western Georgia. During excavations in 2007-2010, Middle and Upper Paleolithic layers were identified here. The faunal complex is dominated by the Caucasian turk (Capra caucasica) and the European bison (Bison cf. bonasus). The high proportion of animal bones with traces of cuts and blows means that their accumulation is mainly due to human activity. The characteristics of the complex suggest that the periods of mobile Upper Paleolithic hunter-gatherers ' habitation in the cave were short-lived. The composition of faunal remains suggests that the Upper Paleolithic landscapes around the cave were more open than in the Holocene.

Key words: Late Pleistocene, Caucasus, hunting, paleoecology.

Introduction

Studying the transition from the Middle Paleolithic to the Upper Paleolithic in the South Caucasus is important for understanding the nature of the replacement of Neanderthals by modern anthropological humans in the region where their interaction may have occurred [Adler et al., 2008; Pinhasi et al., 2012; Moncel et al., in press]. Recent research has provided a wealth of data on the behavioral characteristics of ancient humans during this important transition, as well as on the pace of this process. Apparently, and in the North,

And in the South Caucasus, Neanderthals and their accompanying Middle Paleolithic industries disappeared several millennia before the appearance of Upper Paleolithic industries, usually considered as created by modern humans (although there are no anthropological finds directly related to stone artifacts), which does not allow us to speak of a direct interaction between the two human species [Pinhasi et al., 2011; Pinhasi,et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., 2011; Pinhasi, et al., Nioradze, Tushabramishvili et al., 2012]. This gap is also evidenced by the lack of continuity in stone technology, as well as other aspects of material culture, for example, bone products first appear only in the early Upper Paleolithic [Adler et al., 2006; Pleurdeau et al., 2007; Bar-Yosef et al., 2011; Moncel et al., 2013]. At the same time, recent studies have shown that in the territory of Western Georgia, the life support and delivery systems of obsidian by Middle Paleolithic Neanderthals were quite similar to those practiced by anthropologically modern people in the Upper Paleolithic (Adler et al., 2006; Adler and Bar-Oz, 2009; Bar-Oz and Adler, 2005; Le Bourdonnec et al., 2012]. Both populations hunted almost exclusively for ungulates, preferring adult animals, while the development of hunting skills allowed them to intercept Caucasian tours on the paths of their seasonal migrations. The methods of transporting carcasses, butchering them, and eating them for food were almost identical in the late Middle and Upper Paleolithic periods, as evidenced by the study of stratigraphic sequences of key sites, such as the Ortvale Klde and Dzudzuan caves (Adler et al., 2006; Adler and Bar-Oz, 2009; Bar-Oz and Adler, 2005). Bar-Oz et al., 2002; Bar-Oz, Belfer-Cohen, Meshveliani et al., 2008; Bar-Oz, Weissbrod, and Gasparian et al., 2012; Moncel et al., 2013) (for data on the North Caucasus, see Hoffecker and Cleghorn, 2000). These observations are based on a limited set of archaeological sources, and additional Middle and Upper Paleolithic complexes in Western Georgia may allow us to assess how typical these characteristics are of Neanderthal and modern human populations in this region.

Recent excavations in the Bondi Cave (Imereti, Republic of Georgia) yielded faunal materials found in the context of Middle and Upper Paleolithic strata (Tushabramishvili et al., 2012). The results of a detailed zooarchaeological and taphonomic study of this complex provide additional data for studying the Paleolithic of the region [Bar-Oz et al., 2012] and reconstructing hunting and life support systems on the southern borders of the Caucasus ridges in the late Pleistocene.

Parking lot and its surroundings

Bondi Cave is located in the Rioni and Kvirila river basins in the Imereti region and in Northwestern Georgia at an altitude of 477 m above sea level (Figure 1). The landscape of this area is now hilly and wooded, cut by deep gorges. The cave is located on the southern slope of a small valley (Fig. 2), located about 30 m above the Tabagrebi River, 2 - 4 km from the cave sites of Ortvale Klde and Dzudzuana.

In 2007-2010, in the frontal part of the cave, near the modern drip line on an area of 12 ^m2, a thickness of 2.8 m deposits was uncovered, including eight archaeological horizons separated by layers of boulders and rubble formed as a result of large cave-ins (Fig. 3). The rock base has not yet been reached [Tushabramishvili et al., 2012]. Excavations revealed two main cultural complexes: the Middle (final stage) and Upper Paleolithic. Layer VII (Middle Paleolithic, including massive plates, elongated flakes, and evidence of the Levallois technology) is radiocarbon dated in the interval

Figure 1. Location of Bondi Cave and other sites mentioned in the text.

2. Entrance to Bond Cave (a) and view from it (b). July 2011.

3. Plan of excavations in the Bond cave, a - excavations in 2007; b-2009; c-2010.

38 700 - 35 000 bp (Hulu-calibrated values - 43 000 - 40 000 layers V-III (Upper Paleolithic, during the period of their formation, an increase in the proportion of lamellae relative to plates is noted, and a bone assigned to the genus Homo was found in layer V) - 24 600 - 14 000 l. n. (Hulu-calibrated values - 29 000 - 17 000) [Tushabramishvili et al., 2012]. Thus, the Upper Paleolithic layers in Bondi Cave may be partially simultaneous with those of Dzuzuana (layers D - B) and Ortwale Klde (layers 4d and 4c) [Moncel et al., in press].

Animal bones and stone artefacts are found in all archaeological horizons. In total, more than 7,000 stone artefacts were found, about 90 % of them in layers II, IV, and V. In small complexes of the lower layers VII and VIII, thick elongated cleavage products are present, which sharply distinguishes them from the Upper Paleolithic ones, which contain

elongated thin plates [Ibid.]. Evidence of the use of fire was found in all layers in the form of burnt flints and microfragments of coal. The faunal materials of Bondi Cave have already been studied earlier, but the research was limited to the identification of species (Tushabramishvili et al., 2012).

Methods

Zoological and taphonomic data were obtained according to the procedure described earlier [Bar-Oz and Adler, 2005; Bar-Oz et al., 2008]. The entire collection of bones collected during excavations in 2007-2010 was scanned to find identifiable parts of the skeleton. Elements that could not be unambiguously assigned to a particular layer or, at least, to a certain period were not taken into account. The method of identifying parts of the skeleton was maximally expanded and included all detectable and calculable bone fragments. Taxonomic determination was made using a comparative collection and osteological catalogues (Schmid, 1972; Pales and Garcia, 1981). The differentiation of bovine bones (Bos or Bison) was carried out according to the method of S. J. Olsen [Olsen, 1960], in the separation of goat bones (Carpa or Ovis), we followed M. A. Zeder and H. A. Lapham [Zeder and Lapham, 2010]. Identification at the species level was verified on the basis of faunal lists developed by A. Vekua and M. Bukhsianidze, who used a comparative collection from the State Museum of Georgia in Tbilisi. Skeletal elements that could not be assigned to a specific species were grouped according to dimension classes (for example: small mammal, small, medium, or large ungulate).

In order to search for various traces of impact, the bone surface was examined using magnifying lenses with a small resolution (*2.5). Changes caused by humans (traces of cutting, heat exposure and deliberate fragmentation), animals (biting of earthmoving animals, traces of crushing by predator teeth, exposure to gastric juice), and post-exposure (weathering, leaching, crushing due to falling stones or walking on bones, traces of the root system of plants, abrasions) were recorded (according to: [Behrensmeyer, 1978; Fisher, 1995]). Fragments of long tubular bones that showed signs of fracture in ancient times were studied for the morphology of fracture planes. Their outlines and angles were taken into account in order to determine whether the bones were broken in a dry or fresh state (for a detailed description of the types of fractures, see Villa and Mahieu, 1991). The degree of integrity of the circumference of long tubular bones (whole, more or less than half) was also taken into account, which served as a fragmentation index (according to Bunn, 1983). In addition, the age assessment was based on changes in the external surface of the long bones (the degree of fusion of the heads), data on the eruption and degree of wear of the lower jaw teeth (Grant, 1982; Stiner, 1990).

Results

Excavations from 2007 to 2010 at Bondi Cave yielded a medium - sized collection of animal bones, consisting mainly of fragments of long tubular bones of medium and large ungulates, although there are some intact bones and jaws. In total, approximately 5,900 fragments larger than 1 cm were found, including 12 identified specimens (NISPS) in the Middle Paleolithic layers and 277 in the Upper Paleolithic layers. The surface of the bones is mostly well preserved.

In the sediments of both periods, the dominant taxa are the Caucasian turk (Capra caucasica) and the bison, most likely its European variety (Bison cf. bonasus) (Table 1). Other ungulates are represented by single specimens. These include red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Predator remains are present in the sediments of both periods. A wolf (Canis lupus) and a bear (Ursus sp.) were identified. Judging by the relatively small size of the bones, the bear is represented by the species U. arctos, but the small number of currently available measurements does not allow us to state this with certainty (Table 2). A significant part of the collection consists of bones (mainly fragments of diaphysis of long bones), which can only be distributed by size. This classification corresponds to the species ratio: the vast majority of fragments belong to large ungulates (probably bison) and medium-sized ones (the majority here, apparently, are tours, the minority are red deer). The presence of bison (Bos primigenius) is recorded in neighboring sites [Bar-Oz et al., 2008], in contrast to the Bondi Cave complex, where all identifiable ruminant bones show morphological characteristics of bison.

Although the collection from the Middle Paleolithic strata is very small (probably because only the upper part of the Middle Paleolithic deposits has been uncovered so far), we were able to reveal an important difference in the taxonomic composition of the Middle and Upper Paleolithic complexes (Figure 4). The proportion of large carnivore remains (wolf and bear) relative to the proportion of ungulate bones is significantly higher in the Middle Paleolithic layers (χ ² = 10.14, p = 0.001).

Table 1. Taxonomic composition of the collection

Note: NISP is the number of identified bones, MNI is the minimum number of individuals for mutually exclusive species.

Table 2. Bone Measurements

End of Table 2

Notes: measurement method and designations according to (Driesch, 1976); SP - Middle Paleolithic, VP - Upper Paleolithic. * Measurements were made at the junction of enamel and dentin.

4. Structure of Upper and Middle Paleolithic complexes.

a - ungulates; b - large carnivores.

The Upper Paleolithic deposits were divided into three parts: the lower one (layer VI). middle (IV, V) and upper (I-III). Throughout the sequence, the proportion of bony remains of deer and large carnivores is always small, with tur and bison bones dominating in all layers (Figure 5). However, the specific weight of these two taxa varies over time: in the lower and middle parts of the sediments, the first one dominates, and in the upper part, the second one dominates (upper : middle levels - χ = 7.76, p = 0.051).

Due to the small sample size, the taphonomic origin of Middle Paleolithic faunal remains is difficult to determine. One bone with cutting marks (the calcaneus of a deer) and one with a predator's bite (a fragment of the tibia of a medium-sized hoofed animal) were found here. These findings may indicate that both humans and predators participated in the formation of this complex. Much more data are available on the taphonomy of Upper Paleolithic faunal materials (Table 3). The anthropogenic origin of ungulate remains follows from

5. Trends in the distribution of the frequency of occurrence of species in the Upper Paleolithic complex.

Table 3. Data on surface changes and bone fragmentation for each taxonomic group in the Upper Paleolithic complex

* Hereafter, after the slash, the number of instances suitable for defining this feature is indicated.

Fig. 6. Samples with traces of human and animal impact, a-a fragment of the tibia (middle part) of an average ungulate with signs of stone use for its crushing (a wide shell-shaped notch, an internal negative of chipping a chip, a spiral fracture); b - a fragment of the humerus of the tour (N 177), showing the bite of a predator (jagged edges, tooth marks, a narrow notch without traces of internal chipping of the chip).

high specific gravity of bones with cuts and traces of blows. Cuts were found on bones from all parts of the skeletons of tours and bison. They reflect all the stages of cutting the carcass (skinning, cutting, cutting meat, cleaning the periosteum). The nature of the broken bones indicates their crushing with stones. Most tubular limb bones retain less than half of their original circumference, which is often accompanied by a "fresh" character of the fracture angles. This may indicate that the bones were deliberately broken in order to extract their contents (Figure 6 (a)). The almost complete absence of burnt bones (one sample) seems strange. There are no traces of anthropogenic impact on the bone remains of carnivorous animals. The activity of predators in the cave can be attested due to tooth marks on several ungulate bones (Table 3, Fig. 6, b). However, the small number of predator remains and the moderate frequency of rodents suggests that their role in the formation of the Upper Paleolithic faunal complex was small. Influence of abiotic destruction processes (weathering, trampling, impact of the plant root system, abrasions) the impact on bone preservation was negligible (Table 3).

The composition of skeletal elements of medium and large ungulates in the Upper Paleolithic complex was as follows:

It was reconstructed based on the calculation of the minimum number of individuals (MNI) for the most numerous detectable bones in each anatomical department (Table 4). These results can be used only with allowance for bone preservation, which depends, in particular, on bone density. Fragments of diaphysis outnumber epiphysis of the same limb bones of medium and large ungulates (48 vs. 27 and 41 vs. 8, respectively). This means that taphonomic processes affected skeletal elements differently, depending on bone mineral density: the spongy epiphysis was preserved worse than the cortical diaphysis of the same bones.

The results of the analysis of skeletal elements indicate an almost complete representation of the anatomical parts of medium-sized ungulates (tur), while the composition of these parts of large ungulates (bison) looks more selective (Table 4, Fig. 7): mostly lower ones

See Table 4. Representation of skeletal elements of medium and large ungulates in the Upper Paleolithic complex

7. Distribution of skeletal elements of large (a) and medium (b) ungulates in the Upper Paleolithic complex.

jaws and diaphysis of the long bones of the limbs. The low representation of axial skeletal parts in both size classes probably stems from different bone densities, which had a particular impact on the preservation of ribs and vertebrae. It can be assumed that the carcasses of tours were transported in full or almost complete form, while much larger bison were first butchered at the site of extraction, after which the most nutritionally advantageous parts (containing a lot of meat or fat) were transferred to the cave.

The existing collection does not allow us to reconstruct a detailed picture of the age structure of the faunal complex. Based on the analysis of the sequence _of dp4-_M3 teeth with the stage of wear in the Upper Paleolithic collection, it can be assumed that at least two adult tura individuals were consumed. Judging by the stage of teething and the degree of wear of the lower P4, at least two young (non-erased teeth) and two adult (erased teeth) bison were obtained. The degree of fusion of the limb bones indicates the prevalence of adult individuals of tours and bison. The proportion of non-sinostosed bone elements is 14 % for Capra and 20 % for Bison (the total number of detectable individual elements is 35 and 10, respectively). Thus, based on data on teeth (the stage of eruption and the degree of wear) and synostosis of bones, it can be assumed that hunters produced mainly adult tours. Regarding bison, the picture is more vague and contradictory due to the insufficient sample size.

Five processed bones were found in Bondi Cave (Table 5, Figure 8). All of them come from the Upper Paleolithic strata, with the exception of one with an unclear context. Four items are awls or fragments of them. One piece is made from deer antler, the rest are made from long tubular bones of ruminants. All awls, with the exception of-

See Table 5. Samples of processed bones

Notes: VP - Upper Paleolithic, PDTK-fragment of a long tubular bone.

Figure 8. Bone tools from Bond's cave.

a-WB3, awl; b-WB2, awl tip; c-WB4, presumably awl fragment; d-N 160, processed horn tip; e-WB1, presumably awl fragment.

only one of them is fragmented, and none of them bears traces of fire. Items from the Bondi Cave resemble tools of this type from the Dzuzuana Cave [Bar-Yosef et al., 2011, fig. 5, 6], although they are less variable, probably due to a smaller collection.

Discussion

Bondi Cave contains Middle and Upper Paleolithic faunal complexes dominated by tur and bison. This corresponds to stratigraphic sequences in Western Georgia, especially in the nearby Dzudzuana and Ortvale Kld caves [Bar-Oz and Adler, 2005; Adler et al., 2006; Bar-Oz et al., 2008; Adler and Bar-Oz, 2009; Bar-Yosef et al., 2011; Moncel et al., 2013]. The faunal composition is also similar to that found in the Late Middle Paleolithic layers of the Northwestern Caucasus (Hoffecker and Cleghorn, 2000). Our conclusions were based mainly on the Upper Paleolithic collection due to its sufficient size for analysis. These faunal remains indicate that they appeared in the cave as a result of hunting, subsequent transportation and butchering of the extracted animals for meat and bone marrow. Transportation methods depended on the size of the carcasses. Medium-sized hoofed animals (tur) were brought to the camp whole or almost whole, and much larger bison were partially butchered at the site of prey. It is quite possible that large amounts of bison meat were transferred to the campsite in separate chunks (Monahan, 1998). The discarded remains of prey were then eaten by predators (especially wolves). They also underwent the destructive effects of abiotic processes, as a result of which mainly bones characterized by a denser structure were preserved.

The almost complete absence of burnt bones is of interest. At nearby Paleolithic sites, many samples with traces of thermal impact were found [Bar-Oz and Adler, 2005; Bar-Oz et al., 2008], which could be related to the use of pre-existing bone waste. Given this fact and the fact that there is evidence of predator activity in Bondi Cave, as well as a moderate number of bones and individual animals, we are inclined to conclude that the cave was inhabited for short periods by Upper Paleolithic people, who were characterized by high mobility. Taking into account the large proportion of predator remains in the upper Middle Paleolithic layers, it can be assumed that the cave was used even less intensively in the Middle Paleolithic. However, judging by the composition of animal skeletal elements and traces of anthropogenic impact on the bones, some animals were butchered and eaten directly in the parking lot. The almost complete representation of the body parts of the tour suggests that the hunting grounds were located near the cave. Our conclusion is in good agreement with the features of the Middle Paleolithic stone inventory, which is very small and represents an incomplete reduction sequence. This indicates a seasonal short-term occupation of the cave, when ready-made tools were brought. However, the high saturation of the Upper Paleolithic layers with the products of stone splitting may indicate the utilization of local flint in situ and, accordingly, a different character of cave settlement. These data contribute to the discussion about the type of use of the surrounding landscape in the Middle and Upper Paleolithic by the area's populations.

At the end of the Upper Paleolithic stratigraphic sequence, the bison's food use is higher than that of the tura. A decrease in the proportion of bison bone remains during the Upper Paleolithic was also observed in the Dzuzuana Cave, which was perceived as a reflection of changes in the seasonality of settlement of the site [Bar-Oz et al., 2008; Bar-Yosef et al., 2011]. More research is needed to determine whether this trend is really related to seasonality, as well as changes in hunting equipment or the availability of certain animal species.

The two main animal species represented in faunal complexes throughout the entire stratigraphic sequence of Bondi Cave - the Caucasian tur and the European bison - thrive perfectly in the conditions of mountains that combine both forest and open landscapes [Adler et al., 2006; Julien et al., 2012]. The late Pleistocene macrofauna reconstructed from the materials of the monument may indicate lower elevations and more open landscapes than in the Holocene, since the structure of the Mesolithic (Early Holocene) fauna of this region looks different. At the nearby Kotias Klde site, there are no tur and bison bones in the Mesolithic strata, but brown bear, red deer, and roe deer bones dominate, and wild boar (Sus scrofa) is also represented [Meshveliani et al., 2007; Bar-Oz et al., 2009]. The rarity or absence of the last three species in the faunal complexes of Bondi Cave and its contemporary sites probably reflects the conditions of the much more open landscapes of this region in the late Middle and Upper Paleolithic periods. Due to the nature of the faunal complex, we did not notice any noticeable changes in the environment in the Upper Paleolithic. Probably landscape conditions

in the vicinity of the cave and its location on the territory of Western Georgia contributed to the maintenance of life here for most of the late Pleistocene, in contrast to areas with a larger altitudinal area or more remote from the sea (Vekua and Lordkipanidze, 1998; Bar-Oz et al., 2012).

Acknowledgements

We are grateful to A. Vekua and M. Bukhsianidze for their help in taxonomic definitions, and to G. Bar-Oz for their assistance in the course of research and comments on the manuscript of this article. We are also grateful to the French Ministry of Foreign Affairs and the National Museum of Natural History (Paris) for their financial support in fieldwork and analysis.

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The article was submitted to the editorial Board on 20.01.14, in the final version-on 24.01.14.

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