Buccal Dental Microwear and Diet of the Sunghir Upper Paleolithic Modern Humans

ARCHAEOLOGY, ETHNOLOGY & ANTHROPOLOGY OF EURASIA

ELSEVIER

Archaeology Ethnology & Anthropology of Eurasia 42/2 (2014) 131-142 E-mail: [email protected]

ANTHROPOLOGY B. Pinilla1 and E. Trinkaus2 ‘University o f Barcelona, Avenida Diagonal 645, 08028 Barcelona, Spain E-mail: beatriz.pinilla@gmail. com 2Washington University, Saint Louis, MO, 63130, USA E-mail: [email protected]

BUCCAL DENTAL MICRO WEAR AND DIET OF THE SUNGHIR UPPER PALEOLITHIC MODERN HUMANS

Buccal microwear analysis o f the deciduous molars o f Sunghir 3 provides a moderate density o f striations and suggests a mixed diet. The permanent teeth o f Sunghir 1 to 3 present a low density o f microwear, in agreement with the minimal occlusal wear o f Sunghir 2 but in contrast with the more advanced wear o f Sunghir 1 and 3. The Sunghir 1 and 2 scratches imply mixed diets; those o f Sunghir 3 suggest a more carnivorous one. These results are in agreement with bone chemistry and stable isotope data and with the resources likely present near Sunghir. Keywords: Diet, ecology, teeth, Upper Paleolithic, Late Pleistocene, Europe.

Introduction

The site of Sunghir in northern Russia (Bader, 1978) represents one o f the m ore northern early U pper Paleolithic sites in Eurasia and certainly one of the more substantial ones. As a result, it is of interest to assess the paleoecology o f the Sunghir early modem humans to the extent possible, from their geoarchaeological context and from the human remains themselves. The former aspects have been extensively assessed (Bader, 1978, 1998; Alekseyeva, 1998; Gugalinskaya, Alifanov, 2000; Lavrushin, Sulerzhitsky, Spiridonova, 2000), and inferences regarding their dietary profiles have been made from chemical and isotopic analyses (Kozlovskaya, 2000a, b; R ichards et al., 2001; D obrovolskaya, Richards, Trinkaus, 2011). Here we add to this mosaic of information through the analysis o f the dental buccal

microwear of the three Sunghir individuals preserving teeth, Sunghir 1, 2 and 3.

The Sunghir site and human remains

The Paleolithic site of Sunghir is adjacent to Vladimir City (56°10’30" N; 40°30'30" E), 200 km north of Moscow (Bader, 1978, 1998). Excavated from 1956 to 1977, the site consists of an extended open-air settlement area, probably ~10,000 m2 of which -4500 m2were excavated. The cultural layer is a sequence of paleosols between an underlying sandy-loam and overlying loess levels, but the original levels were extensively mixed by post-depositional solifluction and ice-wedges, obscuring most of the internal stratigraphy (Bader, 1978; Gugalinskaya, Alifanov, 2000). The faunal assemblage (Alekseyeva, 1998) consists of a

Copyright © 2014, Siberian Branch of Russian Academy of Sciences, Institute of Archaeology and Ethnography of the Siberian Branch of the Russian Academy of Sciences. Published by Elsevier B.V. All rights reserved, doi: 10.1016/j.aeae.2015.01.013

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mix o f 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 m ore tem perate species (.Bison sp., Canis lupus, Equus ferus, Gallus sp., Maries martes, Panthera spelaea, Ursus cf. arctos). Associated palynological samples indicate an alternation between pine (Firms), birch (Betula) and spruce (Picea ) dominated woods, but with evidence o f more temperate tree pollen (Lavrushin, Sulerzhitsky, Spiridonova, 2000). This mix o f colder and more temperate faunal and floral species, and especially the presence o f a paleosol within the loess sequence o f the Sunghir area, suggests a m oderately warm period during the Interpleniglacial (MIS 3). It has been referred to the Bryansk Interstadial (Gugalinskaya, Alifanov, 2000) and therefore should correspond to one o f the Greenland Interstadials (GI) (Svensson et al., 2008; Fleitmann et al., 2009). A series o f radiocarbon dates have been determined from the Sunghir faunal rem ains (m ostly m am m oth) (Sulerzhitsky, Pettitt, Bader, 2000; Marom et al., 2012). They range from -2 9 ,5 0 0 to -2 0 ,0 0 0 14C BP, but the majority are between -26,000 and -29,500 14C BP. The date o f -26,000 14C BP is close to the Heinrich Event 3 (HE3) (Hemming, 2004) and therefore does not match the paleoclimatic indicators from the cultural layer. The GI-6 and GI-5 warm peaks are close to 29,500 and 28,00014C BP, respectively (Svensson et al., 2008), suggesting that the Sunghir cultural layer dates to one o f these Greenland Interstadials. The most likely interstadial is GI-5, given the cluster o f dates closer to 28,000-27,000 l4C BP; the post-HE3 GI-4 is too recent for the radiocarbon dates. T he S unghir 1 to 3 hum an rem ains derive from spectacular burials (Sunghir 1 in Grave 1 and Sunghir 2 and 3 in Grave 2) that were dug into the sandy-loam sediments below the cultural layer (Bader, 1998). There have been m ultiple attem pts to directly date the hum an rem ains

(Pettitt, Bader, 2000; Kuzmin et al., 2004; Dobrovolskaya, R ichards, Trinkaus, 2011; M arom et al., 2012), and questions have arisen as to their contemporaneity with each other and the cultural layer, due largely to more recent dates for Sunghir 1. However, there was no indication of the burials being dug through the cultural layer, a culturally similar burial (Grave 2bis) was within the cultural layer, and similar artifacts are known from both contexts (Bader, 1978,1998). Moreover, the more reliable direct dates, those from (Kuzmin et al., 2004) for Grave 2 and (Dobrovolskaya, Richards, Trinkaus, 2011) for Graves 1 and 2, place them -27,000 to -2 6 ,0 0 0 14C BP. Given paleoclimatic indicators for correlation with an interstadial (and hence unlikely to have occurred during HE3) and association with the cultural layer, they probably derive from GI-5. Sunghir 1 to 3 consist o f largely complete skeletons, with damage principally to the axial remains o f Sunghir 1 and loss o f the left distal upper limb o f Sunghir 2. The first is an adult male, with an age-at-death o f 35—45 years, Sunghir 2 is an early adolescent (11-13 year-old) male, and Sunghir 3 is a late juvenile (9-11 year-old) probable fem ale (B uzhilova, K ozlovskaya, M ednikova, 2000; M ednikova, Buzhilova, Kozlovskaya, 2000; GuatelliSteinberg, Buzhilova, Trinkaus, 2013; Trinkaus et al., 2014). Sunghir 2 retains a m inim ally worn dentition, with all except the M 3s in occlusion. Sunghir 3 has a dentition in transition, with the Is, Cs, M is plus dm Is and dm2s in occlusion, and the premolars and more distal molars in their crypts or erupting at the time o f death. The Sunghir 1 dentition was extensively worn, such that only the M3s retain most o f the crown, but small amounts of enamel remain on most o f the teeth (Fig. 1).

Microwear methods and samples Methods. To assess the buccal microwear o f the Sunghir teeth, the original teeth were cleaned with acetone, a soft

Fig. 1. Buccal views of the postcanine teeth for Sunghir 1 (SI), 2 (S2), and 3 (S3). The left mandibular dentition of Sunghir 3 is reversed.

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brash and compressed air, and then high resolution molds were taken of the buccal surfaces with Coltene President Jet light-body polyvinylsiloxine. Epoxy (Epo-tek 301) casts were then made from them (cf., (Perez-Perez, Lalueza, Turbon, 1994; Perez-Perez et al., 2003; Galbany, Perez-Perez, 2004; Galbany et al., 2005)). The casts were mounted on stubs and sputter-coated with a 400 A gold layer to allow SEM observation. Only postcanine teeth were analyzed, as anterior dental microwear might be affected by paramasticatory activities (Puech, 1979; Larsen, 1985; Bermudez de Castro, Brom age, Fernandez-Jalvo, 1988; Lalueza, Frayer, 1997; Bax, Ungar, 1999; Lozano et al., 2008). Teeth altered by postmortem damage or artifacts were not included (cf. (King, Andrews, Boz, 1999; Perez-Perez

133

et ah, 2003; M artinez, Galbany, Perez-Perez, 2004; Teaford, 2007)) (Fig. 2, d-f, h). Micrographs (Fig. 2) were taken with Leica 360 (PCB) and Hitachi S3000N (SSTT-UA) Scanning Electron M icroscopes using the standardized m ethodology (Perez-Perez, Lalueza, Turbon, 1994; Perez-Perez et al., 2003; Galbany et al., 2005, 2009). Images were taken at 100* magnification and with a 35^10 mm working distance. Each image was then cut to a 0.56 mm2square area and the scratches were recorded with Sigma Scan V (SPSS) in a semiautomatic procedure considering as scratches all of the objects produced by abrasion on the enamel longer than 15 microns and at least four times longer than wide without considering the curvature (intraobserver error is ~6 %, similar to errors obtained

Fig. 2. SEM images at 100x. a-Sunghir 1 RM 1; b - Sunghir2 LM,; c -S u n g h ir 3 Rdm1; rf-Sunghir 1 LM2; e -S u n g h ir2 LM2;/-S u n g h ir3 L M ,;g -P atau d 1 RP,; h - Rond-du-Barry 1 RM3; i - Pavlov 1 RM2. Images d - f and h represent non-preserved surfaces due to erosion/patina defects (d), micro-pitted surface (e), and abrasion (f, h)\ images a-c and g -i represent surfaces in which scratches are clearly distinct and can be measured.

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in buccal and occlusal analyses (Grine, Ungar, Teaford, 2002; Galbany et al., 2005). Sigma Scan automatically registers the quantity of scratches, their lengths and slopes. Fifteen variables were obtained including the number (N), lengths (X) and standard deviations (S) of the length of the horizontal (0°-22° and 158°-180°, H), vertical (67°-112°, V), mesiodistal (112°-157°, MD), and distomesial (23°-67°, DM) orientations. The three final variables correspond to the total sum of the densities (NT), the mean of the lengths (XT), and the mean standard deviation (ST). A compendium o f the 15 variables defines a buccal microwear pattern (Galbany et al., 2009). The proportion o f horizontal against the vertical scratches has been shown to be a good discriminator between principally carnivorous and primarily vegetarian human populations (Lalueza, Perez-Perez, Turbon, 1996). To avoid over­ representation o f the individuals, the mean o f the micro wear values of the well-preserved teeth is used for each individual’s microwear signature. Although there is a contrast in the microwear of the deciduous versus permanent teeth o f Sunghir 3 (see below), no significant differences were found between the permanent tooth microwear data from Sunghir 3 or other immature specimens in the comparative sample (Les Rois 1 and 2, Brassempouy 884, Cistema 1) and data from mature specimens. Therefore, the data from the permanent teeth of these individuals are included in the analysis with the adults. A K o lm o g o rov-S m irnov one-sam ple test was perform ed using all available teeth to compare the cumulative distributions for the microwear variables with a nonnal distribution. None of the variables is nonnormally distributed (p > 0.05), and therefore parametric tests are applied. Tooth preservation. Although Sunghir 1,2 and 3 retain 20, 16 and 8 postcanine teeth in occlusion respectively, only a subset of each provided reliable buccal microwear. The Sunghir 1 teeth are heavily worn (Fig. 1), three teeth with buccal enamel (the right P4 and the left M2 and M3) are eroded, and the right M2 has only one horizontal scratch. Sunghir 2 retains all of his postcanine teeth, albeit with the M3s in their crypts, but only two teeth (the right P4 and M,) provide microwear. The left P4 to M3 are surface eroded, the left P3 has no scratches, and the right M2 has only one horizontal scratch. Sunghir 3 is missing her left dm1 and dm, (probably through exfoliation) and the right M„ and her M2s are unerupted. Of the remaining teeth, two deciduous teeth (the right dm1and the left dm2) and one permanent molar (the right M1) provide buccal microwear. C om parative sam ples. The Sunghir m icrow ear patterns were compared to a sample o f 35 European Upper Paleolithic individuals, divided between MIS 3 and 2, or Early and Late Upper Paleolithic (Table 1).

The primary sample of comparison is the Early Upper P aleolithic one, given the age and archaeological association of the Sunghir sample. Moreover, given the paleoecological concerns of the analysis, the Early Upper Paleolithic sample is further divided into those from the central Europe (Dolni Vestonice, Mladec, Pavlov, and Predmosti), and those from more the maritime regions of southwestern France (Brassempouy, Isturitz, Pataud, and Les Rois) and northern Italy (Barma Grande and Grottedes-Enfants/Fanciulli). The Late Upper Paleolithic sample derives entirely from southwestern Europe. In addition, distributions of recent human samples, pooled by agricultural versus foraging economies, and by broad climatic categories for the foraging groups (Lalueza, Perez-Perez, Turbon, 1996) were employed. The groups consist of an agricultural sample (central India), tropical forest foragers (Andamanese and Veddah), arid and mesothermal foragers (Native Australians, Tasmanians, and San), and higher-latitude (more “carnivorous”) foragers (Inuit, Saami, Fuegians, and Pacific Northwest Native Americans).

Results The D eciduous M olars o f Sunghir 3. The average microwear pattern of the three Sunghir 3 postcanine teeth differs from those o f the other Sunghir individuals, especially on the density variables (Table 2). One-way ANOVA and Tukey post-hoc analysis indicate that the differences are significant between Sunghir 1 and 3 on the density of the vertical scratches (NV, p = 0.013) and between Sunghir 2 and Sunghir 3 on mesiodistal scratch lengths and the standard deviation o f the scratch lengths (XMD,/> = 0.019; SMD,/> = 0.014). When analyzing the differences in detail, the main distinction is due to the deciduous teeth. Within Sunghir 3, the deciduous molars present a higher density of scratches and shorter ones than the Sunghir 1 to 3 permanent teeth (Fig. 3). The differences between the permanent and deciduous teeth are significant in one-third of the microwear variables (NH, p = 0.020; NV, p = 0.010; XMD, p = 0.029; NT, p = 0.003; XT, p = 0.038). Contrasts between the microwear on deciduous and permanent teeth have been noted previously, and it is probably due to differences in enamel properties (Perez-Perez, Lalueza, Turbon, 1994; Pinilla, Romero, Perez-Perez, 2011) (but see (Gamza, 2010)). Yet, it should be noted that the mean lengths and density of the Sunghir 3 deciduous scratches fall close to the distribution of permanent teeth from other Upper Paleolithic humans (Fig. 4), whereas it is the permanent Sunghir teeth which are outliers from that distribution (see below). Permanent tooth buccal microwear. When the total length (XT) and density (NT) values for the Upper

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B. Pinilla and E. Trinkaus / Archaeology, Ethnology and Anthropology o f Eurasia 42/2 (2014) 131-142

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Table 1 (end)

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B. Pinillci and E. Trinkaus /Archaeology, Ethnology and Anthropology o f Eurasia 42/2 (2014) 131-142

137

200 Paleolithic specimens are compared (Fig. 4), there S2 P, O a is little difference across the Early and Late Upper O Paleolithic samples (p = 0.199), and within the Early □ b 180Upper Paleolithic sample there is a tendency for the S1 M1 O central European ones to have a higher density and 160 shorter lengths of the scratches than the more southern S2 M, ones (p = 0.038). There is nonetheless a general 140S3 M1 inverse relationship between density and length O (r = 0.418). The Sunghir specimens are among the S1 M1 120 higher Upper Paleolithic values in scratch lengths, but □ they are separate from the other specimens in densities. S3 dm2 They are closest to Barma Grande 2, Cisterna 2, 100 □ Ranchot 80, Rond-du-Barry 8, and Grotte-desS3 dm1 Enfants 3; the Upper Paleolithic individuals showing 80 50 100 150 200 250 lower densities of scratches and higher lengths are all Density of scratches per 0.56 mm from more southern portions of Europe. Comparison of the full variable set also indicates Fig. 3. Comparison of the mean length of scratches (XT) versus the that the Sunghir individuals are clearly different from density of scratches (NT) for the individual Sunghir teeth. the European Upper Paleolithic humans. The contrast, - permanent teeth; b - deciduous teeth. as with NT, is due to the lower level of abrasion observed on Sunghir teeth, and it is reflected in all of 180 03 o the density variables (NH,/? = 0.004; NV, p = 0.030; A □ b NMD, p = 0.020; NDM, p = 0.028; NT, p < 0.001). o S2 Oc 160A Principal Component Analysis (PCA) results in A d a similar distinction, with the Sunghir individuals O',S1 differing from the mean of the comparative samples on OS3 A 140□ A Q\ the first component (PCI; 46.96 % of the variability, O mainly related to length variables (XT, r = 0.96; XV, A □ O □ r = 0.90; XH, r = 0.84) and largely on the second □ 120 □ component (PC2; 13.44 % of the variability; mainly O related to density variables: NT, r = 0.57; NH, iO rf? 100 r = 0.55; NDM; r = 0.52). □ 0 □ The Linear Discriminant Analysis (LDA) shows that the Sunghir individuals do not present a common 80pattern with respect to the Upper Paleolithic groups. 50 150 250 350 450 One-way ANOVA shows significant differences Density of scratches per 0.56 mm between the Early and Late Upper Paleolithic samples Fig. 4. Comparison of the mean length of scratches (XT) versus only on vertical scratch density (NV, p = 0.044); the the density of scratches (NT) for the Sunghir and Upper Paleolithic overall LDA correct classification rate is 88.6 %, permanent teeth. 62.9 % with cross-validation (jackknife). The a- Sunghir; b, c - Early Upper Paleolithic of Central (6) and southwestern (c) LDA (non-significant) places Sunghir 1 with the Europe; d - Late Upper Paleolithic. Early Upper Paleolithic sample (56.6 % posterior probability), while Sunghir 2 and 3 are classified as Late Upper Paleolithic individuals (78.3 % and 74.1 % difference between the Upper Paleolithic samples. If the posterior probabilities, respectively). This result is mainly indices are plotted against each other (Fig. 6), there is due to the lengths of the scratches. Function 1 (100 % of an inverse relationship between them across the Upper the variability explained) is mainly represented by NV Paleolithic samples ( r = 0.499) and an intermixing of the (r = 0.39), XDM (r = 0.37), and XT (r = 0.35). Early and Late Upper Paleolithic values. Sunghir 2 and Assessment of the relative proportions of scratches in especially Sunghir 1 are similar to the Upper Paleolithic different orientations provides little difference between pattern in both comparisons (Fig. 5,6), albeit towards the the average Sunghir values and the Upper Paleolithic end of the distribution of indices with a lower proportion samples (Fig. 5). The same similarity is evident in the of vertical scratches and more horizontal ones. They are indices from the proportions of horizontal (NH/NT) among Barma Grande 2, Caldeirao 1, Dolni Vestonice 32, and vertical (NV/NT) scratches to the total number of Grotte-des-Enfants 4, Mladec 1, Pataud 1, Ranchot 80, scratches (Table 3). In these comparisons, there is little and Predmosti 26 and 30. -i

-

138

B. Pinilla and E. Trinkaus/Archaeology, Ethnology and Anthropology o f Eurasia 42/2 (2014) 131—142

Fig. 5. Distributions o f the proportion of horizontal (NH), vertical (NV), mesiodistal (NMD), and distomesial (NDM) scratches for the overall means of the Sunghir, Late Upper Paleolithic, and Early Upper Paleolithic samples and for the Sunghir individual mean values.

■

Sunghir mean

Late Upper Paleolithic

Early Upper Paleolithic

Sunghir 3

Sunghir 2

Sunghir 1

NH

□

NV

□

NMD

□

Sunghir 3, in contrast, is different from the other distributions with a dominance o f mesiodistal scratches (Fig. 5). In the com parison o f the indices, she is in line with the overall distribution but beyond it in term s o f having the highest proportion of vertical scratches (67.0 %) and a low proportion (0.9 %) of horizontal ones (Fig. 6). The closest specimens to it are Les Rois 1 and 2 and Mladec 10. Although the Les Rois specimens are immature - 10-11 years (Vallois, 1958), three other immature individuals, Brassempouy 884 — 10 years (H en ry -G am b ier, M au reille , W hite, 2004), Cistema 1 — 8 years (Trinkaus et al., 2011), and Sunghir 2 -1 1 -1 3 years - are towards the other end of the distribution. Therefore the differences observed in the pattern are not related to the individual’s age-at-death, geography or chronology. N e v erth eless, if the data from all o f the individuals are compared to the indices from the recent hum an sam ples, the U pper P aleolithic individuals range between tropical-arid mixed diets and the more carnivorous hunter-gatherers (cf. Fig. 6,7) (Lalueza, Perez-Perez, Turbon, 1996). Sunghir 1 falls in the overlap zone o f all three

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Horizontal scratch proportion

Table 3. Mean, minimum, and maximum values for the density (NT), length (XT), and indices (NH/NT and NV/NT) of buccal microwear scratches, comparing Sunghir individuals with the Early (EUP) and Late (LUP) Upper Paleolithic samples

E U P (N = 25)

LU P (N = 10)

Sunghir (N = 3)

M ean

NT

XT

N H /N T

N V /N T

2 8 3 .5

124.09

0.17

0 .3 9

Min.

178

94 .57

0.05

0 .1 3

Max.

442

176.35

0.31

0.56

M ean

2 5 3 .3

140.21

0.20

0.32

Min.

185

110.04

0.10

0 .1 3

M ax.

334

171.46

0.33

0.49

M ean

93.2

152.26

0.17

0.44

Min.

82

143.00

0.09

0.32

M ax.

106

166.16

0.25

0.67

B. Pinillci and E. Trinkaus / Archaeology, Ethnology and Anthropology o f Eurasia 42/2 (2014) 131-142

recent hunter-gatherer groups, and Sunghir 2 is close to it (Fig. 7). The implication is that they would have had a mixed diet. In contrast, Sunghir 3 is distant from both of them (as in Fig. 6), in the upper range of the variation of higher latitude foragers subsisting mainly on marine and terrestrial vertebrates.

Discussion The analysis of the buccal microwear of the three Sunghir individuals highlights four aspects. There is a very different pattern between the permanent teeth and the deciduous teeth of Sunghir 3. All three individuals exhibit a low level of dental buccal abrasion in their permanent teeth. Sunghir 1 and 2 have microwear patterns that are common among Early and Late Upper Paleolithic humans, as well as recent human hunter-gatherers; they are distinct principally from the recent agriculturalist sample. Sunghir 3 presents a different microwear pattern, distinct from Sunghir 1 and 2, just beyond the known Upper Paleolithic variation, and at the upper limits in terms of “camivory” for recent foragers. The im p licatio n s o f the buccal m icrow ear o f Sunghir 1 and 2, a mixed vertebrate and plant diet with a low level of abrasiveness, are variably in agreement with the other indicators from their remains and from the site of Sunghir. In terms of abrasiveness, Sunghir 2 has an exceptionally low level of dental wear, even for its early adolescent age-at-death; this low level of attrition may correlate with his minimal masticatory muscle markings (Trinkaus et al., 2014). Sunghir 1, however, exhibits a marked overall level o f occlusal wear, and an exceptional degree of lingual M 1 and M2 wear with a lesser degree of buccal M, and M2 attrition. It is unclear what would have produced the generally low level of buccal abrasiveness; the colder climate may have promoted it, but the sediment underlying the principal cultural layer is a sandy loam. Given the multifactorial origins of these scratches (Puech, Prone, 1979; Peters, 1982; Teaford, Lytle, 1996; Mahoney, 2006; Alrousan, Perez-Perez, 2008), the low density of scratches could be related to differences in processing techniques or other particles ingested with the food. More directly related to diet, Sunghir 1 has yielded trace element proportions that suggest a diet with of a high proportion o f vertebrate meat (Kozlovskaya, 2000a), and stable isotope analysis of both individuals provides 815N values (10.7 % o and 11.2 % o , respectively) th at place them w ell w ithin the carnivore range (R ichards et al., 2001; D obrovolskaya, R ichards, Trinkaus, 2011). The latter values, however, only reflect the individual’s sources of protein, and they do not indicate the proportions o f calories derived from plant versus animal sources. It is likely that they had

139

access to starchy underground tubers (Hardy, 2010), and grindstones from sites further to the south and west indicate that such starchy foods were being processed during the Mid Upper Paleolithic (Revedin et al., 2010). There is also evidence in dental calculus for cooked plant foods among earlier Upper Paleolithic humans in Europe (Henry, 2010). Although the site of Sunghir is much further north, the sedimentology and the mixed ecological indicators (see above) indicate that such resources were likely to have been available at the time of the site’s paleosol formation. It is less straight-forward to account for the different buccal microwear of Sunghir 3. Her dentition shows a normal (for an Upper Paleolithic individual) degree of occlusal abrasion for her age-at-death. The chemical composition of the bone is ambiguous as to diet, with a suggestion of more plant food in the diet (Kozlovskaya, 2000a). Her collagen provides a 815N value of 11.0 % o , between those of Sunghir 1 and 2 and well within the carnivore range (Dobrovolskaya, Richards, Trinkaus, 2011). However, it is not clear why her diet, as suggested by the buccal microwear, should contrast with those o f the other Sunghir individuals. One possibility is that it could be related to her congenital deformities (Buzhilova, 2000; Formicola, Buzhilova, 2004), ones which led to pronounced systemic stress during her decade o f life (Buzhilova, 2005; Guatelli-Steinberg, Buzhilova, Trinkaus, 2011) despite the maintenance of a normal Upper Paleolithic activity level (Mednikova, 2005; Cowgill et al., 2012). Interestingly, if one uses the mean NH/NT versus NV/NT values o f her deciduous teeth (0.480 versus 0.141 (Table 2)), they place her well

Fig. 7. Comparison of the proportion o f vertical scratches (NV/NT) versus horizontal ones (NH/NT) for the Sunghir mean values (SI to S3) versus the distributions of recent human hunter-gatherer and agriculturalist samples.

140

B. Pinilla and E. Trinkaus /Archaeology, Ethnology and Anthropology o f Eurasia 42/2 (2014) 131-142

w ithin the U pper P aleolithic scatter (Fig. 6) and closer to the m iddle o f the recent hum an hunter-gatherer variation (Fig. 7). Yet, it is unclear how m icrow ear on deciduous teeth relates to that on perm anent teeth (see above) and w hether such a com parison is valid.

C on clu sion T h e b u c c a l m ic r o w e a r s ig n a tu r e s o b s e rv e d in th e Sunghir individuals point to a clear difference betw een th e ir d eciduous te e th and p erm an en t den titio n . W hen co m paring th e S unghir v alues w ith th o se from U p p er Paleolithic hum ans across Europe, the results show that they present a low er abrasiveness o f their diet (derived from the low er density o f scratches); it is not clear how th is re la te s to th e ab u n d a n ce o f ab ra siv e p a rtic le s in their diets, since Sunghir 1 and 3 show norm al levels o f occlusal w ear fo r the U pper P aleolithic, but Sunghir 2 exhibits an unusually low level o f attrition for his age. A t the sam e tim e, th e positions o f Sunghir 1 and 2 in the index plot (N H /N T versus N V /N T ) are w ithin the U pper Paleolithic variation, and the Sunghir 3 perm anent tooth (but not her deciduous m olars) is on the m argins o f that distribution. T he Sunghir 1 and 2 buccal m icrow ear also falls w ell w ithin the variation o f recent hum an huntergatherers and im plies a m ixed diet. Sunghir 3 differs from them and appears m ore strictly “carnivorous.”

A ck n o w led g m en ts The analysis of the Sunghir human remains in the Laboratory o f Plastic Anthropology, Russian Academy o f Sciences was undertaken with the permission of the late T.S. Balueva, and the support of N.O. Bader, A.P. Pestryakov, and E.V. Veselovskaya. The Upper Paleolithic comparison sample was obtained thanks to a Leakey Foundation grant to A. Perez-Perez in 2003 and their analysis were funded by the Ministry o f Science and Innovation, Spain, with a predoctoral fellowship to B. Pinilla (AP2006-01274) and a research grant (APP-CGL2007-60802/ BTE) to A. Perez-Perez. SEM images were taken at the Serveis Cientificotecnics (UB-PCB) o f the University of Barcelona and at the scanning microscopy unit (SSTT-UA) at the University of Alicante (A. Romero). This analysis is part o f a larger re­ analysis of the Sunghir human remains, in collaboration with A.P. Buzhilova, M.V. Dobrovolskaya, and M.B. Mednikova, and funded by the Russian Foundation for Basic Research and Washington University.

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Alrousan M., Perez-Perez A. 2008 Non-occlusal microwear of the last hunter-gatherers from the Near East and Europe. In Genes, Ambiente y Enfermedades en los Poblaciones Humanas, A.J. Nogues, S. Pinilla (eds.). Zaragoza: Univ. of Zaragoza Press, pp. 45-59. Bader O.N. 1978 Sungir - verkhnepaleoliticheskaya stoyanka. Moscow: Nauka. Bader O.N. 1998 Sungir: Paleoliticheskiye pogrebeniya. In In Pozdnepaleoliticheskoye poseleniye Sungir (pogrebeniya i okruzhayuschaya sreda), N.O. Bader (ed.). Moscow: Nauchny mir, pp. 5-160. Bax J.S., Ungar P.S. 1999 Incisor labial surface wear striations in modem humans and their implications for handedness in Middle and Late Pleistocene hominids. International Journal o f Osteoarchaeology, vol. 9: 189-198. Bermudez de Castro J.M., Bromage T.G., Fernandez-Jalvo Y. 1988 Buccal striations on fossil human anterior teeth: Evidence of handedness in the Middle and early Upper Pleistocene. Journal o f Human Evolution, vol. 17: 403-412. Buzhilova A.P. 2000 A naliz anom alii i indikatorov fizicheskogo stressa u nepolovozrelykh sungirtsev. In Homo sungirensis: Verkhnepaleoliticheskii chelovek: Ekologicheskiye i evolyutsionnye aspekty issledovaniya, T.I. Alekseyeva, N.O. Bader (eds.). Moscow: Nauchny mir, pp. 302-315. Buzhilova A.P. 2005 The environm ent and health condition o f the U pper Palaeolithic Sunghir people of Russia. Journal o f Physiological Anthropology and Applied Human Sciences, vol. 24: 413-418. Buzhilova A.P., Kozlovskaya M.V., Mednikova M.B. 2000 Opredeleniye pola i vozrasta sungirskikh individuumov: Sungir 1. In Homo sungirensis: Verkhnepaleoliticheskii chelo­ vek: Ekologicheskiye i evolyutsionnye aspekty issledovaniya, T.I. Alekseyeva, N.O. Bader (eds.). Moscow: Nauchny mir, pp. 54-56, 62. Cowgill L.W., Mednikova M.B., Buzhilova A.P., Trinkaus E. 2012 The Sunghir 3 U pper Paleolithic juvenile: Pathology and persistence in the Paleolithic. International Journal o f Osteoarchaeology. DOI: 10.1002/oa.2273 Dobrovolskaya M., Richards M.P., Trinkaus E. 2012 Direct radiocarbon dates for the Mid Upper Paleolithic (eastern Gravettian) burials from Sunghir, Russia, Bulletins et Memoires de la Societe d ’Antropologie de Paris, vol. 24: 96-102. Fleitmann D., Cheng H., Badertscher S., Edwards RL., Mudelsee M., Goktiirk O.M., Fankhauser A., Pickering R., Raible C.C., Matter A., Kramers J., Tiiysiiz O. 2009 Timing and climatic impact o f Greenland interstadials recorded in stalagmites from northern Turkey. Geophysical R esearch L etters, vol. 36: L19707. D O I:1 0 .1029/2009 GL040050 Formicola V., Buzhilova A.P. 2004 Double child burial from Sunghir (Russia): Pathology and inferences for Upper Paleolithic funerary practices. American Journal o f Physical Anthropology, vol. 124: 189-198.

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