One of the most important achievements in the Paleolithic archaeology of Eurasia is the proof of the existence of the musteroid culture in the Northwestern Altai. Isotopic studies of bone collagen from late West European Neanderthals have repeatedly confirmed their extreme carnivory. There is practically no information about the feeding habits of Altai Neanderthals. Therefore, we conducted isotope studies to determine the main food resources of the inhabitants of Okladnikov Cave. From seven samples of well-preserved bone tissue of adults and children, collagen was isolated and the ratios of ^13C / ^12C and ^15N/14N were determined. The values of δ13 S indicate the inclusion of all individuals in the local food chain based on plants of the temperate climate zone. The values of Δ15n clearly indicate a high trophic level of these individuals. Based on the results obtained, it can be concluded that the lifestyle of Neanderthals of the Northwestern Altai and late European Neanderthals was similar. Both of them probably specialized in hunting large herbivores.

Key words: Neanderthals, Altai, stable carbon and nitrogen isotopes, paleoecology, paleodietological reconstructions.

Altai Neanderthals: typical features and local specifics

Large-scale systematic studies of the Paleolithic of the Northwestern Altai made it possible to radically reconsider the mechanisms of the formation of modern humanity. The discovery of the early Upper Paleolithic industry of anatomically modern humans and the identification of new forms whose genesis is associated with the preservation of archaic Homo genes through the hybridization of various groups of fossils and anatomically modern humans put the region in the focus of a wide range of problems of human evolution. One of the most important achievements in this area is the proof of the existence of a musteroid culture in the Northwestern Altai (Derevyanko and Markin, 1992; Derevyanko, 2007; Derevyanko, Markin, and Zykin, 2008). Decoding of the mitochondrial genome from bone fragments from Okladnikov Cave, as well as morphological study of skeletal remains, allow us to speak with complete confidence about the settlement of this region by Neanderthals in the late Middle Paleolithic (Turner, 1990; Krause et al., 2007; Buzhilova, 2011; Mednikova, 2011). Recall that these discoveries of the Neanderthal ecumene were immediately expanded by 2 thousand km to the east.

The study was carried out within the framework of the Russian Foundation for Basic Research projects "Environment and dietary traditions of Paleolithic humans based on isotope analysis" (N 10 - 06 - 00447-a) and " Transformation of biological indicators of Pleistocene Homo in the context of Eurasian migrations "(N 11 - 06 - 12009-ofi-m-2011).

After the publication of the article by I. Krause and his colleagues, the question of the presence of Neanderthals as carriers of the Sibiryachikha musteroid culutra in Southern Siberia was resolved. However, the problem of their existence in the more southern parts of Central Asia is still debatable. M. Glantz, referring to previous studies [Glantz, Viola. Chikisheva, 2004], writes that it is impossible to confirm with certainty that individuals from the Obi-Rakhmat and Teshik-Tash caves have morphological features typical of Neanderthals. Rather, according to the author, they are the result of mixing local Homo representatives with migrants from more northern or eastern regions (Glantz, 2010). All the more intriguing are the evidence-based conclusions about the presence of Neanderthals in Southern Siberia.

In recent years and months, new paleogenetic data and concepts have been introduced into scientific circulation, considering the spatial and temporal structure of the existence of Neanderthals in Eurasia. Analysis of the variability of the hypervariative part of the mitochondrial DNA of 12 individuals allowed a team from the Mediterranean University of Marseille to test several scenarios for the formation of local variants within Neanderthal groups [Fabre, Condemi, Degioanni, 2009]. The sample included individuals dated from skeletal remains in the range of 100-29 KA BP, who lived on a vast territory from Western Spain (El Sidron) to the Altai (Okladnikov Cave). Most of them (with the exception of Skladina, Belgium) are dated no earlier than 50 thousand years ago, so the systematization of data refers to the late Neanderthals.

The researchers concluded that there are three groups: western, eastern and southern. The eastern group includes the inhabitants of Teshik-Tash and Okladnikov Caves, while the southern group includes individuals from the eastern part of the Iberian Peninsula, Italy, and the western Balkans (Monte Lessini, Vindia, and El Sidron?). Note that the individual from the Mezmay cave (Caucasus) is among the Western Neanderthals. According to the authors, the three groups identified did not exist in isolation; there were contacts between them. These findings echo the results of another paleogenetic study.

The recently completed work of an international team of scientists from Spain, Sweden, Denmark ,and the United States (Dalen et al., 2012) identifies two subgroups of Neanderthals. The first group includes individuals from Skladina (Belgium, ca. 100 thousand years ago), Teshik-Tash (ca. 70 thousand years ago), Valdegoba (Northern Spain, ca. 48 thousand years ago), Okladnikov Cave (Northwestern Altai, ca. 40 thousand years ago), Mezmay Cave (Caucasus, ca. 36 thousand years AGO) and Monte Lessini (Italy, ca. 50 thousand years ago); in the second - three from Vindia (Croatia, from 42 to 38 thousand years AGO), two each from Feldhofer (France, ca. 39 thousand years ago) and El Sidron (Northern Spain, 49-35 thousand years AGO). According to the authors, some dramatic events occurred around 50 thousand years AGO, which led to a sharp decline in the number of Neanderthals in general, after which only a small population with a low level of genetic variability remained in Europe. In the next few thousand years, before meeting anatomically modern humans, the European population was represented only by descendants of this consolidated group. According to the researchers ' conclusions, the genetic diversity of earlier Neanderthals was large and quite comparable to the genetic variability within modern humanity, but after the dramatic events of about 50 thousand years AGO, it sharply decreased. As follows from the materials of the mentioned monuments, this reduction affected the European part of the Neanderthal ecumene and did not affect the Asian one. Unfortunately, the poor preservation of the DNA of Middle Eastern forms makes it much more difficult to create an overall picture of the dynamics of Neanderthal genetic diversity.

In connection with the presented hypothesis of geneticists, we recall that the appearance of Neanderthal groups in the Northwestern Altai is considered by researchers of this region as a relatively short-term and local phenomenon.: "About 50 thousand years AGO, a small population of Neanderthals with a musteroid industry penetrated the territory of Gorny Altai. Despite the relatively well-studied territory, only two localities are known so far - Okladnikov Caves and Chagyrskaya Caves, where culture - bearing horizons with the Sibyarichikha musteroid industry have been identified" [2011, p.49]. Thus, it can be assumed that the period of about 50 thousand years AGO was a boundary that contributed to the formation of local variability within the Neanderthal continuum. The nature of the events that occurred at this time, which are important for Neanderthals, is not yet clear. However, a wide range of methods for studying the paleoanthropological source allows us to look for ways to solve this problem today. One of them is the study of the lifestyle, adaptation and natural environment of various temporary and territorial groups of Neanderthals.

Paleoecological approaches to the study of Neanderthal dispersal

Traditionally, the morphological features of Neanderthals were considered as adaptive, developed by scientists. -

data obtained over the period of habitation in European landscapes that changed as the weather cooled. In recent years, even these familiar ideas have undergone certain changes. For example, computer tomography and the creation of a three-dimensional model of the nasal sinuses have shown that their relative size in Neanderthals does not exceed that of modern humans [Rae, Koppe, Stringer, 2011]. It follows from this that the Neanderthal's efficiency of warming the air during respiration is no greater than that of the average modern European.

A review article by T. Weaver [Weaver, 2009] also criticizes the accepted views on the adaptive nature of typical features of cranial and postcranial morphology. The author believes that gene drift, rather than adaptation to harsh climatic conditions, played a leading role in the formation of the morphological identity of Neanderthals. T. Weaver justifies his position by referring to the accepted lists of typical Neanderthaloid traits, most of which clearly do not have an adaptive nature. In addition, according to the author, some lifestyle features (type of diet and specifics of physical activity) can also be considered as factors that influenced the morphology of Neanderthals. Nevertheless, such striking morphological features as a relative shortening of the distal limb segments, combined with a significant shoulder width and a massive chest, fully correspond to the Arctic adaptive type described by T. I. Alekseeva [1977]. Therefore, it seems wrong to completely deviate from the concept of the severity of morphophysiological adaptations in Neanderthals. Perhaps we should formulate the reasons for these adaptations more correctly, because for the most part, Neanderthal groups lived in moderately cold climatic conditions that are not comparable to modern Arctic and subarctic ones. In general, the morphophysiological status, which provides greater physical strength, a significant lung volume, and a relatively low level of heat loss, has a number of advantages not only in overcoming low-temperature stress. Other typical features of Neanderthals associated with internal hypermassivity are not really explained by the Arctic adaptive type features.

Note that researchers have repeatedly emphasized the specific hormonal status of Neanderthals, which provides an increased mass of the muscular component and a significant internal massiveness of the skeleton. Based on a number of models, it is assumed that the mechanics of Neanderthal movements are significantly more energy-intensive, along with very high physical loads. This is consistent with the hypothesis of increased body needs for food and, consequently, a much greater dependence on a sufficient amount of it.

In this case, we can expect that markers of physiological stress, presumably associated with malnutrition, will occur more often in Neanderthals than in anatomically modern humans. To test this hypothesis, we can effectively use data on the frequency of occurrence of a marker of physiological stress - linear enamel hypoplasia (LEH).

M. Ogilvie, B. Curran, and E. Trinkaus (Ogilvie, Curran, and Trinkaus, 1989) studied 669 crowns of European Neanderthal teeth and determined the incidence of LEG - 36.0 %. The sample included both baby teeth and permanent teeth. For the former, this figure is only 3.9 %, and for the latter -41.9%. When calculated for the proportion of individuals with this defect, it turned out that 75 % had at least one tooth with LEG. Most rarely, LEH was found on the first molars - teeth with the earliest terms of crown formation. Based on this study, we can draw several conclusions::

1) Neanderthals were generally more susceptible to food stress than the" average " anatomically modern human;

2) biological or social protection that guarantees the growth and development of the body during the formation of baby teeth, as well as the first molar, provided adequate nutrition for children in Neanderthal communities;

3) starting from the later stage of childhood, the body was much less protected. It should be noted that the growth and maturation rates of Neanderthals were higher, so it is rather difficult to judge with confidence the age of transition from "childish" nutrition to "adult" nutrition. It is definitely possible to say that this occurred later than the formation of the crown of the first molar (by modern standards, from about 4.5 to 6 years).

Studies conducted with odontological collections of Neanderthals and near-modern hunter-gatherers (Eskimos from Point Hope, Alaska; representatives of the early stage of the Jomon culture, Japan) allow us to clarify the age of transition to "adult" nutrition. It should be noted that Neanderthals are closest to traditional hunters of high latitudes in terms of their lifestyle and diet. The Neanderthal group included 18 individuals (from Krapina and Southern France), while the Eskimo group included 21 individuals. The frequency of linear enamel hypoplasia in terms of individuals was 38.9% in the first sample, and 37.5% in the second sample [Temple, Nakatsukasa, McGroarty, 2012].]-

The results obtained earlier were based on the materials of the Ekwen and Uelen ancient Eskimo burial sites (Kozlovskaya, 2001), as well as the combined Eskimo series of Alaska (Keenleyside, 1998). Note only that in the Ekwen group, the incidence of LEG in children is significantly higher than in adults. This significantly distinguishes the hunters of the Middle Paleolithic from those close to modern times.

Based on these facts, the previously proposed hypothesis of increased food stress pressure on Neanderthal populations seems to be losing its universality. However, a comparison of the details of the formation of enamel hypoplasia lines in the groups of Neanderthals and modern hunters showed significant differences. For example, Neanderthals were found to have a higher incidence of LEG on incisors compared to Inuit individuals (Temple, Nakatsukasa, McGroarty, 2012). When calculating the perikimat lines included in this defect, it turned out that the duration of stress in Neanderthals did not exceed 85 days (with an average of about 30 days), and Inuit individuals had defects that lasted more than 125 days (with an average of about 60 days). Thus, we can assume that food stress, so typical for all hunters of the past, had a certain peculiarity in Neanderthal groups. The beginning of its active effect occurred at the age of formation of the crown of the upper incisors. It is also important to note that there are no significant differences in the incidence of LEG in adult and immature Neanderthals. So, in general, the data obtained support the assumption that food stress in Neanderthals was not so significant.

Thus, on the one hand, due to the peculiarity of their morphofunctional status, Neanderthals needed a large amount of high - calorie and protein-rich food, and on the other hand, the specifics of the formation of enamel hypoplasia suggest that they did not experience significant periods of hunger or malnutrition. This situation can occur if there is a very good supply of food resources.

Isotopic studies of bone tissue collagen from late West European Neanderthals (Bocherens et al., 2005) have repeatedly confirmed their extreme carnivory. However, due to the poor preservation of organic compounds in warm climates, we did not have systematic information about the nutritional characteristics of Neanderthals in the Levant and the Middle East.

Recently, as part of an international collaboration between universities in Germany, Australia, Austria, and the United States, wear on the crowns of Neanderthal and early Homo sapiens teeth in Europe and the Middle East was studied using optical three-dimensional topometry [Fiorenza et al., 2011]. Scientists came to the conclusion that the ecological and geographical factor was the leading factor in the choice of food strategy for both of them. In the Mediterranean region, they used various food resources, and in higher latitudes they specialized in hunting.

There is practically no information about the dietary features of the Altai Neanderthals based on the study of an anthropological source. Therefore, we set out to conduct isotope studies to determine the main food resources of the inhabitants of Okladnikov Cave. Good preservation of the organic component of skeletal fragments made it possible to carry out paleodietological reconstruction.

Materials and methods

Seven samples of well-preserved dense bone tissue were collected from Okladnikov's cave: the talus, calcaneus, and humerus fragments of adult females*, the patella of an adult male, and two femoral and humerus bones of juvenile individuals. Collagen was isolated from all the samples.

Collagen extraction was performed in the V. N. Sukachev Laboratory of Biogeocenology and Historical Ecology of the A. N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences using the accepted method [DeNiro and Epstein, 1981; Jorkov, Heinemeier, Lynnerap, 2007]. The bone fragment was placed in a monomolar hydrochloric acid solution at +3 ° C until complete demineralization. Then it was washed with distilled water by centrifugation until neutral pH values were obtained. The organic residue was converted to a soluble form at pH 2.5 and 1 70 °C for 24 hours. The solution was lyophilized without filtration. A Thermo-Finnigan Delta V Plus mass spectrometer with an element analyzer (Thermo Flash 1112) was used. The atomic ratio of carbon to nitrogen was 3.2, which is in the accepted range of 2.9-3.6 and confirms the good preservation of bone collagen.

Earlier, samples were taken from two fragments (the humerus bones of an adult woman and a child) for radiocarbon dating. The date of 24,260 ±180 bp (KIA-27010) was obtained for the female bone, and three dates were obtained for the child bone: 29,990 ± 500 (KIA - 27011), 34,860 ± 360 (Beta-186881), and 37,800 ±

*Here and further, we use the definitions of gender and age by M. B. Mednikova [2011].

See Table 7. ¹³ S / ¹² S and ¹⁵ N/¹⁴ N in the bone collagen of the humerus bones of an adult woman and a child from Okladnikov Cave (comparison of the results of four laboratories)

* Data from [Krause et al., 2007].

± 450 (OX-15481) bp [Krause et al, 2007]. Obviously, such significant discrepancies in the results indicate methodological problems in sample preparation*. In addition, no studies have been conducted to assess the effect of cave water carbonates. Therefore, it is still too early to be sure about the age of paleoanthropological materials from Okladnikov Cave.

Our data on stable nitrogen and carbon isotopes partially coincide with the results of the Beta laboratories (Miami) and Kiel University and significantly differ from the Oxford ones (Table 1). How significant are the discrepancies? Probably, only in one case are they significant: the nitrogen isotope index of the child sample obtained at Oxford is more than 1 % higher than our definition, which may affect the interpretation. In all other cases, the difference does not go beyond the usual variation in the isotopic composition of the fossil material.

What are the possible reasons for discrepancies? The sample preparation protocols used by laboratories vary slightly. The developed ultrafiltration technique [Higham, Jacoby, Bronk Ramsey, 2006] can significantly correct the results of significant temporary transformation of collagen.

Results

As follows from the table below. 2 values of δ13 S, the local food chain is based on plants of the temperate climate zone. The Δ15n values clearly indicate a high trophic level of these individuals [Dobrovolskaya and Tiunov, ²⁰¹¹]. Note that the intra-group variability is very small (Figure 1). Even if some samples belong to the same skeleton, the minimum possible number of individuals is three. This similarity of indicators indicates a common type of nutrition for adults and adolescents, men and women. The sample from a fragment of the humerus of an adult female differs slightly in the ratio of ¹³ S/¹² S. It should be recalled that morphologically this fragment "shows similarity in structure with the most gracile forms of the Middle and Upper Paleolithic" (Mednikova, 2011)and is dated much later. Perhaps this individual existed in a slightly different landscape environment.

Table 2. ¹³ S / ¹² S and ¹⁵ N/¹⁴ N in bone tissue collagen of individuals from Okladnikov cave

1. Ratios of ¹³ S / ¹² S and ¹⁵ N/¹⁴ N in bone tissue collagen of individuals (children and adults) from Okladnikov Cave.

* In recent years, radiocarbon dating of Paleolithic sites has undergone significant changes due to technical innovations. This is the application of ultrafiltration of the collagen fraction of the sample [Higham, Jacoby, Bronk Ramsey, 2006] and obtaining dates for a specific amino acid that is part of bone collagen [Chavan et al., 2012].

Reconstruction of the main commercial animal species

To determine the isotopic characteristics of the main commercial animal species that this group of Neanderthals hunted, it is enough to perform a simple procedure for subtracting the value of the enrichment coefficient. The difficulty lies in the fact that it is not constant and varies significantly. For example, for some Upper Paleolithic ecosystems, the wolf-herbivore enrichment coefficient is 0.9-1.3 for carbon and 4.4 - 5.5 for nitrogen (Bocherens and Drucker, 2003). The authors recommend using the intervals 0 - 2% o for ^5-13 S and 3 - 5% o for δ15 ^{N. Rather than using a specific value.} However, such a wide interval significantly reduces the content of our reconstructions.

According to the recommendations, the isotopic parameters of the collagen of animals hunted by Neanderthals who lived in Okladnikov Cave were approximately as follows: δ13 C from -21.4 to -19.4%, δ15 N from 8.7 to 10.7 %. Unfortunately, the faunal remains from the Okladnikov Cave have not been studied, so we will turn to materials originating from the Chagyr Cave, which also contains the presence of carriers of Mousterian traditions. We isolated collagen from bone samples of a herbivore (bison) and a predator (wolf)* and the ratios of ¹³ S/¹² S and ¹⁵ N/¹⁴ N were determined [Tiunov and Dobrovolskaya, 2011]. As shown in Figure 2, the carbon isotope parameters of the proposed production correspond to bison collagen. However, the value of Δ15n (7.1%) obtained for the bone tissue of this animal is slightly lower than the expected value. The high nitrogen content of human collagen could be attributed to the Neanderthals ' regular consumption of river fish, but in this case the carbon content should have been significantly lower. Therefore, this hypothesis is currently unconvincing.

The specific nature of the Neanderthal hunting strategy has been repeatedly discussed. For a more detailed analysis, we can refer to the isotopic parameters of Neanderthal bone tissue collagen from various European sites. Currently, samples from skeletal remains of 14 adult individuals have been studied (according to [Richards and Trinkaus, 2009; Richards, Taylor, Stelle et al., 2008]). The generalized data indicate that the living conditions of Neanderthal groups were quite similar, as well as the strategies for selecting the main commercial species, as evidenced by stable nitrogen indicators (Table 3).

2. Ratios of ^13C / ^12C and ^15N/14N in ^the collagen of human bone tissue (five samples) and animals from Okladnikov and Chagyrskaya caves, a-human; b - bison; c-wolf.

Table 3. Carbon and nitrogen isotopes in the bone collagen of European Neanderthals (according to [Richards and Trinkaus, 2009; Richards, Taylor, Stelle et al., 2008])

The exception is made by individuals from Feldhofer, the northernmost monument. They probably lived in slightly wetter boreal conditions and hunted mainly medium-sized herbivores. Most other groups obviously preferred more open landscapes (forest-steppe, alternation of open and wooded areas). The main prey of European Neanderthals were animals with δ15n ^approx. 7 - 8 ‰. Extensive summaries of the ¹³ C / ¹² C H ¹⁵ N/¹⁴ N ratios in the collagen of various animals of the Late Mousterian period in Europe

* Species definitions of animals were made by Prof. A. K. Aghajanyan.

They will allow us to assess the local and epochal dynamics of the ecological features of their habitat.

As can be seen from the data in Table 4, the individual variability of isotopic parameters for different species is not the same. It depends on the degree of mobility of the species ' way of life, the scale of migrations of herbivorous mammals, and the variety of animal diets. Based on the summary data, individual variability is highest for reindeer and bison. Reindeer make regular long migrations, eat grass, lichens and small animals for food. The ⁸⁷ Sr / ⁸⁶ Sr ratio in the enamel of deer teeth from the layers of the Jonzac monument (Richards, Taylor, Stelle et al., 2008) indicates that these animals killed by Neanderthals while hunting near the site made much longer migrations during their lives than bison killed by the same hunters. It should be noted, however, that the Pleistocene bison is also characterized by long migrations in open sedimentary areas - the high individual variability of the ^5-15 N values can only be a consequence of this (Table 5). The same can be said about the wild horse.

As was shown earlier (Ambrose and De Niro, 1986; Yakumin, Nikolaev, and Ramigny, 2000), the intensity of nitrogen isotope enrichment is higher in arid landscapes. Therefore, we have every reason to assume that the increased values of ^5-15 N obtained for Neanderthals from the Okladnikov Cave are due to the natural and climatic specifics of the environment, as well as the hunting of animals migrating long distances through arid settled spaces. The study of archaeozoological materials from Pleistocene layers of cave sites in Northwestern Altai suggests that the vast majority of mammalian species lived in sedimentary open landscapes (Prirodnaya Sreda..., 2003).

Another factor that could lead to an increase in the values of isotopic indicators is the significant proportion of young animals in the hunting prey of Neanderthals. The isotopic composition of bone tissue collagen in young herbivorous animals during lactation should differ significantly from that of adults. Relative to the mother, the cub occupies the next trophic level and has higher values of δ15 N. Archaeozoological studies in the region also confirm a significant proportion of young animals among the bone remains of animals [Ibid.]. However, the role of this factor should not be overestimated, since this relatively easy prey was available to Pleistocene hunters only in the warm season. Nevertheless, the combination of these two factors could have led to the formation of a peculiar isotopic composition of Neanderthal bone tissue from the Okladnikov Cave.

See Table 4. Limits of individual variability δ13 S, δ15 N for the main commercial animal species based on the Saint-César, La Berbie, Camiac monuments, Southwestern France (according to [Richards, Taylor, Stelle et al., 2008])

Limits of individual variability Table 5. Δ13C, Δ15N for the main commercial animal species based on the Jonzac monument, Southwestern France (according to [Richards, Taylor, Stelle et al., 2008])

Problems of reconstruction of migration features of Neanderthal groups

This raises the question of the degree of mobility of Neanderthal hunters. Our results indicate a uniform diet for adults of both sexes and children. We also have data on a high degree of similarity of individual isotopic parameters for such famous European monuments as Feldhofer, Vindia, etc. This indicates that all the hunting grounds used by the group were located in the immediate vicinity of the settlement site. If individual hunters (or small groups of them) systematically carried out long-term raids, this would be reflected in the specific isotopic composition of their skeleton. Judging by the low individual variability of isotopic parameters in the groups, the prey was equally distributed among the tribesmen. This is possible when the group is sedentary or migrates together.

In recent years, new methodological techniques have emerged that allow reconstructing the mobility of specific representatives of Neanderthals during their lifetime. So, a group of researchers from the Max Planck Institute for Evolutionary Anthropology and the Speleological Center of Greece studied the enamel and dentin layers of the third molar of a Neanderthal and animal teeth from the Lakonis Cave in southern Greece. The ⁸⁷ Sr/⁸⁶ Sr ratios were determined. The data obtained by laser ablation suggested that the area in which the Neanderthal grew up and the one where he lived in adulthood were at least 20 km apart [Richards, Yarvati, Grimes et al., 2008]. However, this author's interpretation caused a discussion. It was suggested that the dynamics of the ⁸⁷ Sr/⁸⁶ Sr ratio over the course of life may also be related to the consumption of various marine products (Nowell and Horstwood, 2009). So, either this individual did not make significant transitions during the formation of the crown of the third molar, or there was a single transition of several tens of kilometers in length.

Mapping strontium concentrations and 87sr^/86sr ratios in the enamel and dentin of the first molar of a three-year-old Neanderthal child from Paire ⁽France) [Gran et al., 2011] using laser ablation also yielded important results. A comparison of the indicators obtained for the tooth and the surrounding geological deposits showed that the child had changed its habitat shortly before death. The nearest area in which the child could have lived before is located 50 km from the place of his death.

Studies of the isotopic composition of the teeth of deer hunted by Neanderthals living at the Jonzac site (France) suggest that this occurred during seasonal migrations of animals through territories close to the site (Britton et al., 2011).

So, the above facts give grounds to assume that migrations were common among Neanderthals, but not frequent, they were a one-time transition for several tens of kilometers, followed by a long period of sedentary life. These very general conclusions are, in our opinion, extremely important for understanding the lifestyle of Neanderthals. Those groups that are available to us for a comprehensive study undoubtedly belong to specialized hunters (leaving aside the options for southern adaptive strategies). The main commercial animals for them were large herbivores with varying degrees of seasonal migration. The information we know about the Neanderthal lifestyle suggests that they could not have made systematic long-term migrations (following herds), so the choice of place of residence was of paramount importance. Near this area, there should have been migration routes for herds, as well as animals that were constant prey. Recall that guaranteed food supply, and in large quantities, is one of the indispensable living conditions of Neanderthals. Therefore, areas where it was possible to find shelter (caves in low mountains), food (open spaces rich in grassy vegetation), a constant source of fresh water, became the place of their stationary habitat. These are the characteristics of the areas where the Okladnikov and Chagyrskaya caves are located. The ⁸⁷ Sr/⁸⁶ Sr ratio in the enamel of the Okladnikov Cave tooth (Latkoczy et al., 2004) indicates that this individual spent his life in this area without leaving it. Probably, the exceptionally favorable conditions found by a group of Neanderthals who came to the Northwestern Altai allowed them to lead a sedentary lifestyle for some time. However, the fact that representatives of other Homo species may have lived here at the same time suggests that the struggle for possession of such an attractive territory could have resulted in deterioration of hunting conditions or extermination for Neanderthals. For their successful existence, first of all, it was necessary to own hunting grounds. The presence of competitors or changes in natural conditions should have led either to migration in search of new hunting grounds, or to the destruction of competitors. Taking into account all the advantages of-

Given the above facts and considerations, we assume that the presence of a small group of Neanderthals in the territory developed by other Homos could hardly have had a long-term perspective.

As mentioned above, the dramatic events are ca. 50 thousand years AGO led to a sharp decline in the number and diversity of the European Neanderthal population. The groups that lived later in Europe were descended from a small, genetically homogeneous population. It can be assumed that the absence of representatives of other Homo species here for several millennia made possible a secondary increase in the number of European Neanderthals. Later, the appearance of anatomically modern humans in Europe prevented Neanderthals from using the best hunting grounds as freely, which inevitably led to a decline in their numbers and extinction of their population. In the Northwestern Altai, where the " Denisovans "were competitors of the Neanderthals, the" fate " of the latter could not be as prosperous as in Europe. This is probably why their stay in this area, which is exceptionally favorable in terms of environmental parameters, was a relatively short episode.

In the summer of 2011, new skeletal remains of Neanderthals were found in the Chagyr cave. I hope that they will help us understand the genesis of this group in more detail. According to the results of genetic studies by L. Dalen and his colleagues, the carriers of the Mousterian traditions of the Northwestern Altai are descendants of the early Eurasian population of Neanderthals. Studying the morphology of postcranial remains from the Okladnikov Cave allowed M. B. Mednikova to conclude that "the South Siberian and Near Asian forms from the Okladnikov and Tabun caves are specifically similar due to their supposed origin from the general population of early Neanderthals" [2011, p.86].

How genetically homogeneous was the Altai group? In the case of a high degree of its homogeneity, we can speak of a unique migration throw of a small Neanderthal population to the east. If there is genetic variation, it is possible to consider Asian populations as a significant specific share of the Eurasian Neanderthal continuum.

Conclusion

Isotopic studies of Neanderthal skeletal remains from the Okladnikov Cave allowed for a paleodietological reconstruction. The obtained data confirm the hunting specialization of this group. Based on the results obtained, it can be concluded that the lifestyle of Neanderthals of the Northwestern Altai and late European Neanderthals was similar. Both of them specialized in hunting large herbivores. New data allow us to judge the ecological specifics of the existence of the inhabitants of the Okladnikov Cave. The prey that ensured their survival was mainly the fauna of open spaces, which were more arid than the landscapes developed by European Neanderthals. To date, we do not have any facts confirming that the main part of it was made up of the largest herbivores, as it was proved for the Neanderthal groups of Europe. To find out whether the inhabitants of Okladnikov Cave specialized in hunting animals of certain species or made extensive use of local faunal diversity, it is necessary to study in more detail the isotopic indicators for many commercial species. There is every reason to assume that hunting young ungulates was an important component of the hunting strategy of this group.

A number of biological features typical of Neanderthals indicate that the latter are extremely dependent on efficient hunting, which puts them in a vulnerable position when competing for territory with other Homo representatives. This may explain the predominant distribution of Neanderthal hunters in Europe, which is not inhabited by humans, and, obviously, the much less successful history of their development of vast areas of Asia, where evidence of earlier human migrations has been preserved.

Acknowledgements

The study of bone fragments from the Okladnikov and Chagyrskaya caves became possible thanks to the permission of Academician A. P. Derevyanko, Professor M. V. Shunkov, and Professor St. Markin. The authors express their gratitude to the staff of the V. N. Sukachev Laboratory of Biogeocenology and Historical Ecology of the A. I. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences for their assistance in conducting the training.

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