The microstateness of buccal surfaces of permanent teeth in all Sungir individuals is weak, which is consistent with the minimal erasure of occlusal surfaces in individual Sungir 2, but contrasts with their stronger wear and tear in individuals Sungir 1 and 3. The nature of grooves on the teeth of individuals Sungir 1 and 2 indicates a mixed diet. The buccal surfaces of the mammary molars of the Sungir 3 child show a moderate density of wear grooves, which also indicates a mixed diet, but his diet seems to have been dominated by meat. Our results are generally consistent with those obtained in the analysis of the chemical and isotopic composition of bone matter. They are also consistent with data on the natural resources of the area where the parking lot is located.

Key words: diet, ecology, teeth, Upper Paleolithic, Late Pleistocene, Europe.

Introduction

The Sungir site in Central Russia (Bader, 1978) is one of the northernmost and undoubtedly most important Upper Paleolithic sites in Eurasia. Therefore, it would be interesting to study the paleoecology of its inhabitants in as much detail as possible, using both geological and archaeological materials, as well as human remains themselves. Researchers have already turned to geoarchaeological sources more than once [Bader, 1978, 1998; Alekseeva, 1998; Gugalinskaya, Alifanov, 2000; Lavrushin, Sulerzhitsky, and Spiridonova, 2000], and the diet was evaluated based on the chemical and isotopic composition of bone matter [Kozlovskaya, 2000a, b; Richards et al., 2001; Dobrovolskaya, Richards, Trinkaus, 2012]. In this article, an attempt is made to supplement the available, still incomplete information with the results of an analysis of the erasure of the buccal (buccal) surfaces of the teeth of three individuals buried at the Sungir site-Sungir 1-3 (the teeth of other individuals are not available for study).

Sungir site and human skeletons found on it

The Upper Paleolithic Sungir site is located near Vladimir (56° 10 ' 30 "N, 40° 30' 30 " E), 200 km north of Moscow (Bader, 1978, 1998). Excavations were carried out in 1956-1977. It is a large open-type monument with an area of perhaps about 10,000 ^m2. The excavated area is 4,500 ^m2. Cultural deposits consist of several layers of buried soils lying on sandy loam and overlain by loess. Cultural horizons were so severely disturbed by solifluction and ice veins that stratigraphy could not be traced in most areas (Bader, 1978; Gugalinskaya and Alifanov, 2000). The faunal complex (Alekseeva, 1998) is represented by cold-adapted species (Dicrostonyx cf. torquatus. Gulo gulo, Lagarus cf. lagarus, Lepus timidus, Lyrurus tetrix, Mammuthus primigenius, Microtus sp., Ocotona sp., Rangifer tarandus, Saiga cf. tatarica, Spermophilus citellus, Vulpes lagopus) and to temperate climates

(Bison sp., Canis lupus, Equus ferus, Gallus sp., Martes martes, Panthera spelaea, Ursus cf. arctos). Spore-pollen analysis revealed several alternating phases with a predominance of different tree species in the forest cover-pine (Pinus), birch (Betula), and spruce (Picea), but in general, trees adapted to a temperate climate dominated [Lavrushin, Sulerzhitsky, and Spiridonova, 2000]. A mixture of cold-tolerant fauna and flora with those that have been adapted to a more temperate climate, and especially the presence of buried soil within loess deposits in the area of the monument indicate a moderately warm interstadial within MIS 3. It was named the Bryansk interstadial (Gugalinskaya and Alifanov, 2000) and probably corresponds to one of the Greenland interstadials (Svensson et al., 2008; Fleitmann et al., 2009).

A series of radiocarbon dates were obtained from animal bones (mainly mammoths) from the Sungir site (Sulerzhitsky, Petit, and Bader, 2000; Marom et al., 2012) in the range of 29.5 - 20.0 Ka BP, and most of them are in the range of 29.5-26.0 Ka BP.This is the result of the Heinrich event (Hemming, 2004) and thus contradicts the paleoclimatic parameters obtained from the materials from the cultural layer. The dates of warm Greenland interstadials 6 and 5 (GI-6 and GI-5) are 29.5 and 28.0 Ka BP, respectively (Svensson et al., 2008). The Sungir cultural layer probably dates back to one of them, most likely GI-5, whose date is closer to the concentration of radiocarbon dating sites-28 - 27 KA. b. Dates of the Greenland interstadial 4, following the 3rd Heinrich event, later than the Sungir ones.

Skeletons of Sungir 1-3 were found in the richest graves (Sungir 1 - in grave 1, Sungir 2 and 3 - in grave 2), sunk into the sandy loam layer that underlies cultural deposits (Bader, 1998). Several attempts have been made to directly date human remains (Pettitt and Bader, 2000; Kuzmin et al., 2004; Dobrovolskaya, Richards, and Trinkaus, 2012; Marom et al., 2012). The date of the Sungir 1 skeleton turned out to be relatively late, which raised two questions: are these burials simultaneous and how do they relate to the cultural layer of the site? There is no evidence that the graves cut through the cultural layer. The archaeologically similar border 2 was located in the cultural layer, and it did not differ from it in terms of inventory (Bader, 1978, 1998). In addition, more reliable dates for burials 2 (Kuzmin et al., 2004; Dobrovolskaya, Richards, and Trinkaus, 2012) and 1 (Dobrovolskaya, Richards, and Trinkaus, 2012) are close - 27 - 26 Ka BP, taking into account paleoclimatic data, according to which the site dates back to the Bryansk Interglacial and is unlikely to be located in the same area. coincides in time with the 3rd Heinrich event, as well as the connection of burials with the cultural layer, they probably need to be attributed to the time of the Greenland interstadial5.

All three bones are almost complete. Only the axial skeleton of individual Sungir 1 and the distal part of the bones of the left arm of the child Sungir 2 were affected. The skeleton of Sungir 1 belonged to an adult male who died at the age of 35-45 years, Sungir 2 - to a boy of 11-13 years, Sungir 3 - to a child of 9-11 years, possibly a girl [Buzhilova, Kozlovskaya, Mednikova, 2000; Mednikova, Buzhilova, and Kozlovskaya, 2000; Guatelli-Steinberg, Buzhilova, and Trinkaus, 2011; Trinkaus et al. , in prep.]. The teeth of the Sungir 2 boy were very weakly erased, and the upper and lower teeth, except for both M3, were in contact. The dental system of the Sungir 3 girl was in a transitional state, and there was occlusion between the upper and lower I, C, M1, m1 and m2, while the premolars and more distal molars were in holes or erupting at the time of death. The teeth of the Sungir 1 man are almost completely worn out, only the M3 crowns are mostly preserved. However, some amount of enamel is still present on most of his teeth (Figure 1).

1. Posterior teeth of Sungir individuals 1 (S1), 2 (S2) and 3 (S3) in buccal norm. The left lower teeth of individual Sungir 3 are shown in a mirror image.

Erasure assessment methods and material

Methods. To study the erasure of buccal surfaces, the teeth of all individuals were cleaned with acetone, a soft brush and compressed air, and then negative impressions were made from Coltene President Jet (light-body) polyvinylsiloxane mass, according to which casts were made from Epo-tek 301 epoxy mass (see, for example: [Perez-Perez, Lalueza and Turbon, 1994; Perez-Perez et al., 2003; Galbany and Perez-Perez, 2004; Galbany et al., 2005]). They were mounted on pins and coated with a 400-A-thick layer of gold by spraying. After that, the casts were examined by scanning microscopy.

Since anterior teeth wear can be caused by their use in labor operations (Puech, 1979; Larsen, 1985; Bermudez de Castro, Bromage, and Fernandez-Jalvo, 1988; Lalueza and Frayer, 1997; Bax and Ungar, 1999; Lozano et al., 2008), only premolars and molars were studied. Teeth with postmortem injuries and tool marks were not considered (see, for example, [King, Andrews, Boz, 1999; Perez-Perez et al., 2003; Martinez, Galbany, Perez-Perez, 2004; Teaford, 2007]) (Fig.).

Scanning electron microphotographs (Figure 2) were taken using Leica 360 and Hitachi S3000N instruments using standard techniques [Perez-Perez, Lalueza, Turbon, 1994; Perez-Perez et al., 2003; Galbany et al., 2005, 2009]; magnification x 100,

2. Scanning electron microphotographs of teeth at magnification *100, a-Sungir ¹, right M1; b-Sungir 2, left _M1; c-Sungir 3, right ^M1; d-Sungir 1, left _M2; e-Sungir 2. left ^M2; e-Sungir 3, left M ₁; w-Pato 1, right P ₃; w-Ron du Barry 1, right M ³, i-Pavlov 1, right M ². Images g-g, and show surfaces damaged by erosion or patina (d), minute pits (e), and abrasion (e, i); images a-b, f, and z show surfaces with distinct wear grooves that are available for measurement.

working distance 35-40 mm. Each image is cropped to a square with a side of 0.56 ^mm2. The erasure grooves were recorded using the Sigma Scan V (SPSS) program starting from a length of 15 microns, provided that the length exceeded the width by at least 4 times. The curvature of the grooves was not taken into account. The observation error was approx. 6 %, as in other studies of buccal and occlusal erasure [Grine, Ungar. Teaford, 2002; Galbany et al., 2005].

The Sigma Scan program automatically registers the number of grooves, their length and slope. A total of 15 indicators were obtained, including the number of grooves (N), their length (X), standard deviations (S) of the length of horizontal (oriented at angles 0 - 22° and 158 - 180°, H), vertical (67 - 112°, V), mesio-distal (112 - 157°, MD) and distomesial (23-67°, DM) grooves, as well as the total sum of their location density (NT), average length (XT), and average standard deviation (ST). The combination of these indicators determines the type of buccal erasure [Galbany et al., 2009]. It is known that the ratio of the number of horizontal and vertical grooves is an effective indicator of meat and vegetable diets [Lalueza, Perez-Perez, Turbon, 1996]. To avoid overrepresentation of individuals, average wear rates for well-preserved teeth were calculated for each individual.

Although the milk and permanent teeth were erased to varying degrees in the Sungir 3 girl (see below), the erasure of the latter does not show significant differences from that of other Late Paleolithic children involved for comparison (Leroy 1 and 2, Brassampuy 884, Sisterna 1), as well as in adults. Therefore, data on permanent tooth wear in Sungir children are combined with the corresponding data on adults.

The normality of the accumulated frequency distributions of wear indicators for all the studied teeth was checked using the Kolmogorov-Smirnov criterion. Since no significant deviations from the normal distribution were found in any case (p > 0.05), parametric criteria are applied in the future.

Safety of teeth. Although Sungir individuals 1, 2, and 3 have 20, 16, and 8 occluded posterior teeth, respectively, not all of them have the buccal side available for study. The teeth of the Sungir 1 male are very badly erased (see Fig. 1), right ^P4, left _M2 and _M3, which have preserved enamel on the buccal side, suffered from erosion, and on the right ^M2 there is only one horizontal groove. In the boy Sungir 2, all the posterior teeth (M3 in the alveoli) were preserved, but traces of wear were found only on two - the right _P4 and _M1. The surfaces of the left posterior teeth from ^P4 to ^M3 are eroded, on the left ^P3 there are no grooves, and on the right _M2 there is only one horizontal one. The Sungir 3 girl does not have the left ^M1 and _M1 (perhaps they were lost posthumously), and the right _M1 and both M2s have not yet cut through. Among other teeth, buccal effacement was found on two milk molars (right ^m1 and left _m2) and one permanent molar (right ^M1).

Comparative material. The microextension of the Sungir teeth was compared with that of 35 Upper Paleolithic Europeans. Some of them lived during MIS 3, i.e. in the early period of this epoch, while others lived during MIS 2, i.e. in the late Upper Paleolithic (Table 1). Given the Sungir date, the comparison with the early sample should be considered most significant. Since the paleoecological aspect is particularly important for us, this sample was divided into two subgroups: one from Central Europe (Dolni Vestonice, Mladec, Pavlov, and Przedor), the other from the coast of Southwestern France (Brassampuy, Isturitz, Pato, and Leroy), and from Northern Italy (Barma Grande and Grote Children/Fanciulli). The Late Upper Paleolithic series originates entirely from Southwestern Europe.

In addition, we used the distribution of erasure indicators in two modern combined groups-farmers and hunter-gatherers. The second group was subdivided into regional climatic subgroups (according to [Lalueza, Perez-Perez, Turbon, 1996]). In particular, we used data on farmers of Central India, as well as hunter-gatherers living in tropical forests (Andamans and Veddas), desert areas with temperate climates (Australian Aborigines, Tasmanians and Bushmen) and in high latitudes where the diet is mainly meat-based (Inuit, Sami, fire-Earths and aborigines of the North-West).West coast of North America).

Results

Milk molars of the child Sungir 3. The average indicators of erasure of the girl's back teeth differ from those of other inhabitants of the parking lot. This is especially true for the density parameters (Table 2). Univariate analysis of variance and the Tukey test (post-hoc) indicate that Sungir 1 and 3 significantly differ in the density of the vertical grooves (NV, p = 0.013), and Sungir 2 and 3-in the length of the mesiodistal grooves and its distribution. variability (XMD, p = 0.019; SMD, p = 0.014). A more detailed analysis shows that the differences relate mainly to baby teeth. In the Sungir 3 child, the density of grooves on them is higher, and the grooves themselves are shorter than those on the permanent teeth of all three indies.-

Table 1. Average values of buccal microstrain of teeth in representatives of the Upper Paleolithic population of Europe

End of Table 1

Note. For explanations, see the text.

Table 2. Indicators of buccal microextension on the posterior teeth of individuals from the Sungir parking lot

Notes. Data on teeth with a preserved surface and with traces of abrasion that are available for quantitative study are presented. For explanations, see the text.

3. Average length of grooves (XT) and their density (NT) on individual teeth of Sungir individuals. a-permanent teeth; b - milk teeth.

4. Average length of grooves (XT) and their density (NT) on permanent teeth of Sungirans and other representatives of the Upper Paleolithic population of Europe (a, b, c, Early Upper Paleolithic: Central (b) and Southwestern (c) Europe; d, Late Upper Paleolithic.

3). The differences between permanent and baby teeth are significant for every third indicator of wear (NH, p = 0.020; NV, p = 0.010; XMD, p = 0.029; NT, p = 0.003; XT, p = 0.038). Similar differences have already been noted. Most likely, they are caused by different properties of the enamel of milk and permanent teeth [Perez-Perez, Lalueza, Turbon, 1994; Pinilla, Romero, Perez-Perez, 2011] (see, however, [Gamza, 2010]). It should be noted, however, that the average length and density of grooves on the baby teeth of Sungir 3 children are close to those on the permanent teeth of other representatives of the Upper Paleolithic population of Europe (Fig. 4), while the nature of wear of the permanent teeth of Sungir people is quite different (see below).

Buccal erasure of permanent teeth. The total length of wear grooves (XT) and their density (NT) in the early and late comparative Upper Paleolithic samples are similar (p = 0.199; Figure 4). In the early period, differences between territorial groups are found: the Central European grooves are shorter and their density is higher than in the Southern European ones (p = 0.038). The direction of these differences corresponds to a general negative relationship between the length and density of the grooves (= 0.418). In the Sungir people, the grooves are long, but still fall within the range of variation in the comparative sample, and their density goes beyond these limits. The Sungiris are closest to the individuals from Southern Europe Barma-Yerande 2, Sisterna 2, Ranchot 80, Rhone-du-Barry 8, and Yeroten 3, whose furrows are long and their density is relatively low.

Comparison of the entire set of parameters also shows that the inhabitants of the Sungir site differ significantly from the Upper Paleolithic Europeans. As in the case of NT, the difference is caused by less buccal tooth wear in Sungiris. This is evident in all indicators: NH (p = 0.004), NV (p = 0.030), NMD (p = 0.020), NDM (p = 0.028), and NT (p < 0.001). Principal component analysis reveals that the Sungir population differs from the comparative samples for EKI, which accounts for 46.96 % of the variability. It is correlated mainly with the length of the grooves-XT (r = 0.96), XV (r = 0.90) and XN (r = 0.84). Differences are also found in the ECP, which reflects 13.44 % of the variability and depends mainly on the density indicators-NT (r = 0.57), NH (r = 0.55) and NDM (r = 0.52).

One-factor analysis of variance shows that the differences between the early and late Upper Paleolithic samples reach the level of significance only in the density of vertical grooves (NV, p = 0.044). Discriminant analysis allows us to correctly divide individuals into groups in 88.6 % of cases, and when using the jackknife method-in 62.9 %. The linear discriminant function did not reveal significant differences. The Sungir 1 male is closer to the early Upper Paleolithic group (a posteriori probability of 56.6 %), and the Sungir 2 and 3 children are closer to the late group (78.3 and 74.1%, respectively). This result can be attributed to the length of the grooves; discriminant function 1 (100% variability) is mainly associated with NV (r = 0.39), XDM (r = 0.37) and XT (r = 0.35).

5. The ratio of the number of horizontal (NH), vertical (NV), mesio-distal (NMD), and disto-mesial (NDM) grooves in their total number in Sungirans and representatives of the Early and Late Upper Paleolithic on average and in individual individuals from the Sungir region.

Figure 6. The proportion of vertical (NV/NT) and horizontal (NH/NT) grooves in the Sungir people and other Upper Paleolithic inhabitants of Europe.

Comparison of the proportion of grooves of different orientations does not reveal any significant differences between Sungirans and other Upper Paleolithic people (Fig. The similarity is also found in the proportion of grooves of one orientation or another in the total number of them. This applies to both horizontal (NH/NT) and vertical (NV/NT) grooves (Table 3). If the index values are plotted on the graph (Figure 6), it can be seen that there is a negative relationship between them (= 0.499), and the early and late Upper Paleolithic groups they don't differ. According to these indicators, Sungir 2 and especially Sungir 1 are similar to other people of the Upper Paleolithic era (see Figures 5, 6), although the proportion of vertical grooves in them is relatively low on the overall scale, and the proportion of horizontal grooves is relatively high. In this respect, they are close to the individuals of Barma-Yerande 2, Kaldeiran 1, Dolni-Vestonice 32, Erot Children 4, Mladec 1, Pato 1, Ransho 80 and Przedmosti 26 and 30.

Meanwhile, the Sungir 3 girl differs from other individuals (see Figure 5). The proportion of vertical grooves is the highest (67.0%), and the proportion of horizontal grooves is low (0.9%) (see Figure 6). Leroy 1 and 2, as well as Mladech 10, are the closest to her according to these indicators. Individuals from Leroy are children aged 10-11 years [Vallois, 1958], but the other three children are Brassampui 884 (ca. 10 years [Henry-Gambier, Maureille, White, 2004]), Sisterna 1 (ca. 8 years [Trinkaus et al., 2011]), and Sungir 2 (11 - 13 years old) -are in the opposite part of the variability spectrum. Thus, the marked difference is not related to the age, geographical location,or antiquity of the finds.

If all the Upper Paleolithic individuals are compared with modern humans, it turns out that the former occupy an intermediate position between two groups of hunter-gatherers: desert people with a mixed diet and those who are mainly meat - based (see Figures 6 and 7) [Lalueza, Perez-Perez, Turbon, 1996]. Sungir 1 falls into the zone of overlapping values for these groups, and Sungir 2

Table 3. Average, minimum and maximum values of density (NT), length (XT) and the ratios of NH/NT and NV/NT grooves of buccal microextension in Sungirans and representatives of the Early and Late Upper Paleolithic (RVP and PVP) Europe

Note. For explanations, see the text.

Figure 7. Proportion of vertical (NV/NT) and horizontal (NH/NT) grooves in Sungir individuals (S1-S3) in comparison with the corresponding data on modern hunter-gatherers and farmers.

7). It follows that the diet of the Sungir people was probably mixed. On the contrary, the Sungir 3 female is remote from the rest of the Sungirans (Figure 7) and is in the upper part of the spectrum of variability of indicators characteristic of hunter-gatherers of high latitudes, who feed mainly on the meat of marine and terrestrial vertebrates.

Discussion

Analysis of the microstateness of buccal surfaces of Sungir teeth leads to four main conclusions. The nature of erasure of the milk and permanent teeth of the Sungir 3 girl is very different. In all three Sungiris, the buccal surfaces of the permanent teeth are slightly erased. In this respect, Sungir individuals 1 and 2 are close to early and late Upper Paleolithic Europeans and modern hunter-gatherers, but they differ significantly from farmers. The girl Sungir 3 has a different character of erasure than other Upper Paleolithic Europeans. In this respect, it is close to the upper limit of variation for hunter-gatherers with a meat diet.

The nature of microstrain of the teeth in Sungir individuals 1 and 2 indicates a mixed diet, in which both meat and vegetable products were present, and the food was quite soft. This conclusion is generally consistent with other data obtained from bone and other materials from the Sungir site. As for the obliteration of the occlusal surfaces of the teeth, in the individual Sungir 2, it is exceptionally low, even for a child of this age. A low masticatory load is also indicated by extremely weak development of the attachment points of the masticatory muscles [Trinkaus et al., in prep.]. However, in the Sungir 1 male, occlusal effacement is significant, and if the lingual surfaces of ^m1 and ^M2 are extremely strongly erased, then the wear of the buccal surfaces of _M1 and _M2 is weaker.

What causes the relatively low erasure of the buccal surfaces of the teeth is unclear. Perhaps the cold climate contributed to this in some indirect way? The fact is, however, that the main cultural layer of the parking lot lies on loam. Given that the wear of buccal surfaces can depend on a number of reasons [Puech and Pronet, 1979; Peters, 1982; Teaford and Lytle, 1996; Mahoney, 2006; Alrousan and Perez-Perez, 2008], the specific nature of the abrasive nature of the chewed particles should also be taken into account.

If we turn directly to food, the results of microelement analysis indicate that individual Sungir 1 ate mainly meat from vertebrates [Kozlovskaya, 2000a]. The same was shown by isotope analysis: judging by the Δ15n values of both this man and the boy Sungir 2 (10.7 and 11.2%, respectively), the food was approximately the same as that of carnivorous animals (Richards et al., 2001; Dobrovolskaya, Richards, and Trinkaus, 2012). However, these data only indicate the source of protein, but not the ratio of calories obtained from plant and animal foods.

It is possible that the Sungiri diet included sugary plant tubers (Hardy, 2010). This is indirectly evidenced by the terochniki found on the Middle-Upper Paleolithic sites located south and west of the Sungir River (Revedin et al., 2010). In addition, the results of tartar analysis in Europeans of the early Upper Paleolithic period indicate that their diet included plant foods cooked over a fire (Henry, 2010). Sungir, however, is located much further north, but data on sedimentation and other environmental parameters (see above) also indirectly indicate the availability of such food for the inhabitants of this site.

It is less clear why the nature of tooth wear in the Sungir 3 girl is completely different. Occlusal effacement corresponds to that of Upper Paleolithic children of the same age. The results of chemical analysis of her bone tissue do not give any definite indications on the composition of the diet, but, apparently, the proportion of plant food was quite noticeable [Kozlovskaya, 2000a]. According to the content of the Δ15n isotope in collagen (11.0%), the Sungir ³ skeleton is intermediate

between the skeletons of Sungir 1 and 2, therefore, the diet of this girl was also dominated by meat [Dobrovolskaya, Richards, Trinkaus, 2012]. But then it is not clear how her diet differed from that of the other two individuals, judging by the nature of buccal tooth decay. It was assumed that this girl may have had congenital skeletal abnormalities [Buzhilova, 2000; Formicola, Buzhilova, 2004], which caused systemic stress during the first decade of her life [Buzhilova, 2005; Guatelli-Steinberg, Buzhilova, Trinkaus, 2011], despite the fact that her physical activity was apparently low It is typical for the Upper Paleolithic epoch (Mednikova, 2005; Cowgill et al., 2012).

Interestingly, the ratios of NH/NT and NV/NT on baby teeth (0.480 and 0.141, respectively, see Table 1) are similar to those of normal teeth. 2) Sungir 3 girls fall within the limits of variation in Upper Paleolithic people (see Figure 6). Such values are also typical for modern hunter-gatherers (see Figure 7). However, it is not clear how the microstretchedness of baby teeth correlates with that of permanent teeth (see above) and, accordingly, to what extent such comparisons are permissible.

Conclusion

The microstateness of buccal surfaces of milk and permanent teeth of the inhabitants of the Sungir site is quite different. Comparison with data on other representatives of the Upper Paleolithic era from Europe would seem to indicate that the food of the Sungir people was less abrasive (judging by the less dense arrangement of grooves). However, occlusal tooth effacement in Sungir individuals 1 and 3 is approximately the same as in other Upper Paleolithic Europeans, while in Sungir child 2 it is significantly less than in children of this age. The NH/NT and NV/NT ratios of Sungir 1 and 2 individuals fall within the limits of the variation series of these indicators for other Upper Paleolithic European residents, and the Sungir 3 child occupies the extreme place in this series by the nature of wear of permanent (but not milk) teeth. Individuals of Sungir 1 and 2 are similar to hunter-gatherers in terms of the microsternity of buccal surfaces, and their diet seems to have been mixed. Apparently, the diet of the Sungir 3 girl was dominated by meat.

Acknowledgements

The study of the Sungir remains stored in the plastic reconstruction laboratory of the Institute of Ethnology and Anthropology of the Russian Academy of Sciences was carried out with the permission of the late T. S. Baluyeva with the support of I. O. Bader, A. P. Pestryakov and E. V. Veselovskaya. The study of the dental system of other Upper Paleolithic Europeans was made possible by a grant received by A. Perez-Perez from the Lika Foundation in 2003. The laboratory processing of these data was funded by the Spanish Ministry of Science and Innovation through a scholarship for the preparation of the doctoral dissertation of B. Pinilla (project AR2006-01274) and a research grant to A. Perez-Perez (project APP-CGL2007-60802/BTE). Scanning electron micrographs were obtained in the Scientific and Technical Department of the University of Barcelona and in the Department of Scanning Electron Microscopy of the University of Alicante (A. Romero). This study is part of a project to reanalyze Sungir skeletons in collaboration with A. P. Buzhilova, M. V. Dobrovolskaya, and M. B. Mednikova, with financial support from the Russian Foundation for Basic Research and Washington University.

List of literature

Pozdnepaleoliticheskoe poselenie Sungir ' (pogrebeniya i okruzhayushchaya sreda) [Late Paleolithic settlement of Sungir (burials and environment)], ed. by N. O. Bader, Moscow: Nauch. mir, 1998, pp. 240-257.

Bader O. N. Sungir-Upper Paleolithic site. Moscow: Nauka Publ., 1978, 254 p. (in Russian)

Bader O. N. Sungir: Paleolithic burials / / Late Paleolithic settlement of Sungir (burials and environment) / ed. by N. O. Bader, Moscow: Nauch. mir, 1998, pp. 5-160.

Buzhilova A. P. Analiz anomaliy i indikatorov fizicheskogo stressa u nepolovozrelyykh sungirtsev [Analysis of anomalies and indicators of physical stress in immature Sungirians]. Homo sungirensis: Verkhnepaleoliticheskiy chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research].

Buzhilova A. P., Kozlovskaya M. V., Mednikova M. B. Opredelenie pola i vozrasta sungirskikh individuumov: Sungir ' 1 [Determining the sex and age of Sungir individuals: Sungir 1]. Homo sungirensis: Verkhnepaleoliticheskiy chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research]. 62.

Gugalinskaya L. A., Alifanov V. M. Osobennosti pochvoobrazovatel'nogo protsessa po materialam poseleniya Sungir ' [Features of the soil-forming process based on the materials of the Sungir settlement]. Homo sungirensis: Verkhnepaleoliticheskiy chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research].

Kozlovskaya M. V. Rezul'taty khimicheskogo analiza kostnoi tkani podrostkov Sungir 2 i Sungir 3 [Results of chemical analysis of bone tissue in adolescents Sungir 2 and Sungir 3]. Homo sungirensis: Verkhnepaleoliticheskiy chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research].

Kozlovskaya M. V. Sostav mineral'noi chasti kostnoi tkani individua Sungir 1 [Composition of the mineral part of the bone tissue of the individual Sungir 1]. Homo sungirensis: Verkhnepaleoliticheskii chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research].

Lavrushin Yu. A., Sulerzhitsky L. D., Spiridonova E. A. The age of the Sungir I archaeological site

Homo sungirensis: Verkhnepaleoliticheskiy chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [Features of the natural environment of the time of habitation of a primitive person]. Ed. by T. I. Alekseeva, N. O. Bader, Moscow: Nauch. mir, 2000, pp. 35-42.

Mednikova M. B., Buzhilova A. P., Kozlovskaya M. V. Sungir 2 and Sungir 3: sex and age diagnostics according to morphological criteria of the bone system / / Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research / ed. by T. I. Alekseeva, N. O. Bader. - Moscow: Nauch. mir, 2000. - p. 53 - 64.

Sulerzhitsky L. D., Petit P. B., Bader N. O. Radiocarbon age of the settlement and discovered burials of II Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research / ed. by T. I. Alekseev, N. O. Bader, Moscow: Nauch. mir, 2000, pp. 30-34.

Alrousan M., Perez-Perez A. Non-occlusal microwear of the last hunter-gatherers from the Near East and Europe // Genes, Ambiente у Enfermedades en los Poblaciones Huma-nas / eds. A.J. Nogues, S. Pinilla. - Zaragoza: University of Zaragoza Press, 2008. - P. 45 - 59.

Ungar PS. Bax J.S., Incisor labial surface wear striations in modern humans and their implications for handedness in Middle and Late Pleistocene hominids // Int. J. of Osteoar-chaeology. - 1999. - Vol. 9. - P. 189 - 198.

Bermudez de Castro J.M., Bromage T.G., Fernandez-Jalvo Y. Buccal striations on fossil human anterior teeth: evidence of handedness in the Middle and early Upper Pleistocene // J. of Human Evolution. - 1988. - Vol. 17. - P. 403 - 412.

Buzhilova A.P. The environment and health condition of the Upper Palaeolithic Sunghir people of Russia // J. of Physiological Anthropology and Applied Human Sciences. -2005. - Vol. 24. - P. 413 - 418.

Cowgill L.W., Mednikova M.B., Buzhilova A.P., Trinkaus E. The Sunghir 3 Upper Paleolithic juvenile: Pathology and persistence in the Paleolithic // Int. J. of Osteoarchaeology. -2012. -Doi: 10.1002/oa.2273.

Dobrovolskaya M., Richards M.P., Trinkaus E. Direct radiocarbon dates for the Mid Upper Paleolithic (eastern Gravettian) burials from Sunghir, Russia // Bull, et Memoires de la Societe d'Antropologie de Paris. - 2012. - Vol. 24, N 1/2. - P. 96 - 102.

Fleitmann D., Cheng H., Badertscher S., Edwards R.L., Mudelsee M., Gokturk O.M., Fankhauser A., Pickering R., Raible C.C., Matter A., Kramers J., Tuysuz O. Timing and climatic impact of Greenland interstadials recorded in stalagmites from northern Turkey // Geophysical Research Letters. -2009. -Vol. 36: L19707. - Doi:10.1029/2009GL040050.

Formicola V., Buzhilova A.P. Double child burial from Sunghir (Russia): Pathology and inferences for Upper Paleolithic funerary practices // Am. J. of Physical Anthropology. -2004. - Vol. 124. - P. 189 - 198.

Galbany J., Estebaranz F., Martinez L.M., Perez-Perez A. Buccal dental microwear variability in extant African Hominoidea primates: taxonomy versus ecology // Primates. - 2009. - Vol. 50. - P. 221 - 230.

Galbany J., Martinez L.M., Lopez-Amor H.M., Espurz V., Romero A., De Juan J., Perez-Perez A. Error rates in buccal-dental microwear quantification using scanning electron microscopy // Scanning. - 2005. - Vol. 27. - P. 23 - 29.

Galbany J., Perez-Perez A. Buccal enamel microwear variability in Cercopithecoidea primates as a reflection of dietary habits in forested and open savanna environments // Anthropologic. - 2004. - Vol. 42. - P. 13 - 19.

Gamza T.R. Intra-individual microwear variation: deciduous versus permanent dentition // Dental Anthropology. - 2010. -Vol. 23. -P. 66 - 68.

Grine F.E., Ungar P.S., Teaford M.E. Error rates in dental microwear quantification using scanning electron microscopy // Scanning. - 2002. - Vol. 24. - P. 144 - 153.

Guatelli-Steinberg D., Buzhilova A.P., Trinkaus E. Developmental stress and survival among the Mid Upper Paleolithic Sunghir children: Dental enamel hypoplasias of Sunghir 2 and 3 // Int. J. of Osteoarchaeology. - 2011. - Doi: 10.1002/oa.l263.

Hardy B. Climatic variability and plant food distribution in Pleistocene Europe: Implications for Neanderthal diet and subsistence // Quaternary Science Review. - 2010. -Vol. 29. - P. 662 - 679.

Hemming S.R. Heinrich Events: Massive Late Pleistocene detritus layers of the North Atlantic and the global climate imprint // Review of Geophysics. - 2004. - Vol. 42: RG1005. -Doi: 10.1029/2003RG000128.

Henry A.G. Plant foods and the Dietary Ecology of Neandertals and Modern Humans: Ph.D. Thesis / George Washington University. - Washington, 2010. - 284 p.

Henry-Gambier D., Maureille B., White R. Vestiges humains des niveaux de l'Aurignacien ancien du site de Brassempouy (Landes) // Bull, et Memoires de la Societe d'Anthropologie de Paris. - 2004. - Vol. 16. - P. 49 - 87.

King T., Andrews P., Boz B. Effect of taphonomic processes on dental microwear // Am. J. of Physical Anthropology. -1999. - Vol. 108. - P. 359 - 373.

Kuzmin Y.V., Burr G.C., Jull A.T.J., Sulerzhitsky L.D. AMS ¹⁴C age of the Upper Palaeolithic skeletons from Sungir site, Central Russian Plain // Nuclear Instruments and Methods in Physics Research. Ser. B. - 2004. - Vol. 223/224. -P. 731 - 734.

Lalueza C., Frayer D. W. Non-dietary marks in the anterior dentition of the Krapina Neanderthals // Int. J. of Osteoarchaeology. - 1997. - Vol. 7. - P. 133 - 149.

Lalueza C., Perez-Perez A., Turbon D. Dietary inferences through buccal microwear analysis of Middle and Upper Pleistocene human fossils // Am. J. of Physical Anthropology. - 1996. -Vol. 100. - P. 367 - 387.

Larsen C.S. Dental modifications and tool use in the Western Great Basin // Am. J. of Physical Anthropology. -1985. - Vol. 67. - P. 393 - 402.

Lozano M., Bermudez de Castro J.M., Carbonell E., Arsuaga J.L. Non-masticatory uses of anterior teeth of Sima de los Huesos individuals (Sierra de Atapuerca, Spain) // J. of Human Evolution. - 2008. - Vol. 55(4). - P. 713 - 728.

Mahoney P. Intertooth and interfacet dental microwear variation in an archaeological sample of modern humans from the Jordan Valley // Am. J. of Physical Anthropology. -2006. - Vol. 129. - P. 39 - 44.

Marom A., McCullagh J.S.O., Higham T.F.G., Sinitsyn A.A., Hedges R.E.M. Single amino acid radiocarbon dating of Upper Paleolithic modern humans // Proceedings of the National Academy of Sciences USA. - 2012. - Vol. 109. -P. 6878 - 6881.

Martinez L.M., Galbany J., Perez-Perez A. Paleodemography and dental microwear of Homo habilis from east Africa // Anthropologic. - 2004. - Vol. 42. - P. 53 - 58.

Mednikova M.B. Adaptive biological trends in the European Upper Palaeolithic: the case of the Sunghir remains // J. of Physiological Anthropology and Applied Human Sciences. - 2005. - Vol. 24. - P. 425 - 431.

Perez-Perez A., Espurz V., Bermudez de Castro J.M., de Lumley M.A., Turbon D. Non-occlusal dental microwear variability in a sample of Middle and Late Pleistocene human populations from Europe and the Near East // J. of Human Evolution. - 2003. - Vol. 44. - P. 497 - 513.

Perez-Perez A., Lalueza C., Turbon D. Intra individual and intra group variability of buccal tooth striation pattern // Am. J. of Physical Anthropology. - 1994. - Vol. 94. - P. 175 - 187.

Peters C. Electron-optical microscope study of incipient dental microdamage from experimental seed and bone crushing // Am. J. of Physical Anthropology. - 1982. - Vol. 57. -P. 283 - 301.

Pettitt P.B., Bader N.O. Direct AMS radiocarbon dates for the Sungir Mid Upper Palaeolithic burials // Antiquity. -2000. - Vol. 74. - P. 269 - 270.

Pinilla B., Romero A, Perez-Perez A. Age-related variability in buccal dental microwear in Middle and Upper Pleistocene human populations // Anthropological Review. -2011. - Vol. 74. - P. 25 - 37.

Puech P.F. The diet of early man: evidence from abrasion of teeth and tools // Current Anthropology. - 1979. - Vol. 20. -P. 590 - 592.

Puech R.E., Prone A. Reproduction experimental des processus d'usure dentaire par abrasion: implications pa-leoecologiques chez l'homme fossile // Comptes rendus de l'Academie des Sciences. Ser. D. - 1979. - Vol. 289. -P. 895 - 898.

Revedin A., Aranguren B., Becattini R., Longo L., Marconi E., Lippi M.M., Skakun N., Sinitsyn A., Spiridonova E., Svoboda J. Thirty thousand-year-old evidence of plant food processing // Proceedings of the National Academy of Sciences USA. - 2010. - Vol. 107. - P. 18815 - 18819.

Richards M.P., Pettitt P.B., Stiner M.C., Trinkaus E. Stable isotope evidence for increasing dietary breadth in the European mid-Upper Paleolithic // Proceedings of the National Academy of Sciences USA. - 2001. - Vol. 98. - P. 6528 - 6532.

Svensson A., Andersen K.K., Bigler M., Clausen H.B., Dahl-Jensen D., Davies S.M., Johnsen S.J., Muschler R., Parrenin F., Rasmussen S.O., Rothlisberger R., Seierstad I., Steffensen J.P., Vinther B.M. A 60 000 year Greenland stratigraphic ice core chronology // Climate of the Past Discussions. - 2008. - Vol. 4. - P. 47 - 57.

Teaford M.F. What do we know and not know about diet and enamel structure? // Evolution of the Human Diet: The Known, the Unknown and the Unknowable / ed. by PS. Ungar. -N. Y: Oxford University Press, 2007. - P. 56 - 76.

Teaford M.F., Lytle J. Diet-induced changes in rates of human tooth microwear: A case study involving stone-ground maize // Am. J. of Physical Anthropology. - 1996. -Vol. 100. - P. 143 - 147.

Trinkaus E., Bailey S.E., Davis S.J.M., Zilhao J. The Magdalenian human remains from the Galeria da Cisterna (Almonda karstic system, Torres Novas, Portugal) and their archeological context // O Arqueologo Portugues. Ser. V -2011. - Vol. 1. - P. 395 - 413.

Trinkaus E., Buzhilova A.P., Mednikova M.B., Dobrovolskaya M.V. The Paleobiology of the Sunghir People // Etudes et Recherches Archeologiques de l'Universite de Liege (in prep.).

Vallois H.V. Le gisement aurignacien des Rois a Mouthiers (Charente): Les restes humains // Gallia Supplement. -1958. - Vol. 9. - P. 118 - 137.

The article was submitted to the Editorial Board on 10.02.13, in the final version-on 21.02.14.

Permanent link to this publication: