Iron Age Nomads of Southern Siberia in Craniofacial Perspective

Home , Evenks, Pazyryk burials, Prehistoric Mongolia, South Central Siberia, Tagar culture, Ukok Plateau

Anthropological Science Vol. 122(3), 137–148, 2014

Iron Age nomads of southern Siberia in craniofacial perspective

Ryan W. Schmidt1,2*, Andrej A. Evteev3 1University of Montana, Department of Anthropology, Missoula, MT 59812, USA 2Kitasato University, School of Medicine, Department of Anatomy, Sagamihara, Kanagawa 252-0374, Japan 3Anuchin Research Institute and Museum of Anthropology, Lomonosov Moscow State University, Moscow, 125009, Mokhovaya St., 11, Russia

Received 9 April 2014; accepted 24 July 2014

Abstract This study quantifies the population history of Iron Age nomads of southern Siberia by ana- lyzing craniofacial diversity among contemporaneous Bronze and Iron Age (7th–2nd centuries BC) groups and compares them to a larger geographic sample of modern Siberian and Central Asian populations. In our analyses, we focus on peoples of the Tagar and Pazyryk cultures, and Iron Age peoples of the Tuva region. Twenty-six cranial landmarks of the vault and facial skeleton were analyzed on a total of 461 ancient and modern individuals using geometric morphometric techniques. Male and female crania were separated to assess potential sex-biased migration patterns. We explore southern Siberian population history by including Turkic-speaking peoples, a Xiongnu Iron Age sample from Mongolia, and a Bronze Age sample from Xinjiang. Results show that male Pazyryk cluster closer to Iron Age Tuvans, while Pazyryk females are more isolated. Conversely, Tagar males seem more isolated, while Tagar females cluster amongst an Early Iron Age southern Siberian sample. When additional modern Siberian samples are included, Tagar and Pazyryk males cluster more closely with each other than females, sug- gesting possible sex-biased migration amongst different Siberian groups. This is evident in modern female Tuva, who cluster with modern female Kalmyk, while modern Tuvan males do not. Male and female Iron Age Tuvans are not closely related to modern Tuvan peoples living in the region today, resulting from the influx of the Xiongnu beginning in the Late Iron Age. Both male and female Pazyryk and Tagar crania appear more similar to Central Asia groups, especially the Kazakh and Uzbek samples. However, there is evidence that Tagar females have a common origin with the Yakut, a modern nomadic population that resides in northeastern Siberia. These results would suggest variable genetic contribu- tions for both sexes from Central and East Asia.

Key words: craniofacial variation, Pazyryk, Tagar, Tuva, geometric morphometrics

Introduction including canonical variate analysis (CVA) and Mahalano- bis distances between ancient and modern groups from The cultures of southern Siberia, and specifically the southern, western, and eastern Siberia, as well as compara- people of the Altai during the Bronze and Iron Ages, have tive ancient and modern samples from Central Asia, Mongo- previously been studied by physical anthropologists using lia, and western China. traditional craniometric data (Chikisheva, 2000a, b, 2008), The Tagar people are associated with the Late Bronze Age/ however, they have not been examined using geometric Early Iron Age archaeological culture of the same name, morphometrics (GMM). This study investigates two of the which flourished in the Minusinsk Basin along the Upper better-known cultures of the Altai during this time period, Yenisei River from the 7th–2nd centuries BC (Chikisheva, the Tagar and Pazyryk cultures (Figure 1). Although each 2000a). The Minusinsk Basin is located in Khakassia, which cultural tradition has been examined archaeologically, paleo is bordered by the Republic of Altai and the Republic of genetically, and anthropologically, their relationship to one Tuva in the Russian Federation. The Tagar people were another has yet to be fully explored. Here, we present results thought to have been semi-nomadic pastoralists who raised from a craniofacial study that examines population history livestock, such as horses, goats, and sheep. However, based and structure by using exploratory multivariate analyses, on recent archaeological evidence discovered in places such as Mongolia and Kazakhstan, the development of nomadic pastoralism on the eastern steppe was highly complex and * Correspondence to: Ryan W. Schmidt, Ph.D., Kitasato University, School of Medicine, Department of Anatomy, 1-15-1 Kitasato, perhaps included agriculture, in addition to the pastoral Minami-ku, Sagamihara-shi, Kanagawa 252-0374, Japan. economy that epitomizes peoples of the Altai (Di Cosmo, E-mail: [email protected] 1994; Spengler et al., 2014). The production of cereal has Published online 18 November 2014 been speculated among Tagar peoples due to a large number in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.140724 of bronze sickles found in associated archaeological sites

Figure 1. Map showing the extent of of the cultural influence of Pazyryk, Tagar and some adjacent groups in southern Siberia in the 6th–3rd centuries BC.

(Murphy et al., 2013; Svyatko et al., 2013). The Tagar people 2004). AlThough the Scythians may have played a role in also had one of the largest bronze-smelting centers in ancient Tagar development, there is abundant evidence to suggest Eurasia. that peoples of the Tuva region and peoples of the Pazyryk The Pazyryk culture is contemporaneous with the Tagar culture formed via a largely autochthonous component culture, which flourished in the Altai Mountains of southern (Chikisheva, 2008). Siberia and eastern Kazakhstan from the 6th–3rd centuries Therefore, the Altai region has been one of contact, con- BC (Rudenko, 1970). More recently, Pazyryk burial sites flict, and trade since early times. The genetic diversity of the have been found in the Mongolian Altai (Jordana et al., region suggests extensive contact over the course of the last 2009). The Pazyryk people are well known due to the excep- several millennia (Gonzalez-Ruiz et al., 2012). Recent an- tionally well-preserved mummified remains found in stone cient DNA studies indicate the Scythian peoples to have a tumuli of the Ukok Plateau. These burial mounds were in- mix of both Western and Eastern Eurasian DNA types tended for high-ranking members of society, such as chiefs, (Voevoda et al., 2000; Clisson et al., 2002; Ricaut et al., priests, and elders, and included concubines, horse remains, 2004b; Chikisheva et al., 2007; Keyser et al., 2009; Pilipenko and precious artifacts. Archaeologists know that extensive et al., 2010). Nevertheless, the history and origins of these trading networks existed among these nomads as textiles, people remain relatively obscure. Physical anthropologists mirrors, and silk from places such as China, and even Iraq, have suggested the appearance of these peoples, based on have been found among the burials (Chikisheva, 2000a, b). craniofacial traits, to be unlike others in the Altai region Some of the more famous remains include a man and woman during the Bronze and Iron Ages (Rudenko, 1970; Kozintsev with extensive tattooing on their bodies, which depicted et al., 1999; Moiseyev, 2006). Most of the populations resid- elaborate animal motifs. The woman, known as the “Siberi- ing in the Altai during that time possess a more European- an Ice Maiden,” is considered a priestess by archaeologists like appearance relative to populations now residing in the because of the items with which she was buried. region of southern Siberia (Kozintsev, 2009). However, the Some Iron Age nomadic peoples of the Altai, especially Tagar and Pazyryk, when compared to other Bronze Age and Tagar people, have been described as being part of the Early Iron Age samples from the Altai, evidence greater ad- Scythian tradition, though Scythian is a general moniker mixture from Eastern Eurasian populations (Chikisheva, applied to all those tribes who shared economic, cultural, 2000b). Therefore, morphological features more representa- and perhaps linguistic traditions. These mobile equestrian tive of Western Eurasian populations may have appeared as tribes migrated east from the European steppe beginning in early as the Neolithic and the Bronze Age with more recent the 7th century (Ricaut et al., 2004a). Other known tribes admixture occurring during the Late Iron Age in southern that belonged to the Scythian tradition include Sauromatians Siberia (Chikisheva, 2000b). These data have raised ques- of the Lower Volga and southern Urals and the Saka of the tions about the origins and migration routes possibly taken Kazakhstanian steppes and the valleys of Tian Shan and Pa- by the Altai Iron Age nomads. In this study, we use newly mir mountain ranges of Central Asia. The Tagar people pro- acquired craniofacial data analyzed in a GMM framework in duced artwork of animal motifs that were similar in nature to an attempt to tease out the possible origins and interactions works produced by the Scythian tribes. The Scythians are of these Early Iron Age nomads. well known in historical texts and are one of a large number The analysis of ancient populations through the investiga- of tribes to have emerged from the Pontic-Caspian steppe tion of craniofacial variation is an effective and informative and migrated eastward to the Altai from around 2000 BC. way to understand modern population structure and infer The end of the Scythian period was probably the result of the relationships in the past (Howells, 1973, 1989; Hanihara, westward expansion of Turko-Mongolic nomadic peoples 1996). Although morphological variation is subject to devel- from East Asia known as the Xiongnu (Lalueza-Fox et al., opmental and environmental processes that might confound Vol. 122, 2014 ANCIENT NOMADS OF SOUTHERN SIBERIA 139 population history, studies have suggested that the majority of ordinate data were collected using a Microscribe portable cranial traits for populations not residing in extreme climates digitizer by the first author. Male and female adult crania (such as the Arctic) fall under a model of neutral evolution from 201 ancient crania and 260 modern crania are included and are not greatly influenced by selection (Relethford, 2004, in the analyses. All analyses separate sex to assess patterns 2010; Weaver et al., 2008; von Cramon-Taubadel, 2009a; of migration and differential classification. Ancient crania Betti et al., 2010). Therefore, phenotypic variability can be sample names, male and female sample size, site informa- used as a proxy to study biological distances between groups. tion, and periods are displayed in Table 2. Examples of In this study, we use craniofacial diversity as a proxy for Tagar, Iron Age Tuva and Pazyryk male cranial morphology genetic distance between populations. We quantify the dis- are shown in Figure 2. All comparative cranial samples are tance between the ancient male and female Pazyryk and shown in Table 3. Tagar and Pazyryk samples contain indi- Tagar peoples using methods developed in the GMM analy- viduals from several different archaeological sites. In order sis. Although recent studies have used ancient DNA to study to increase sample size, we pooled Tagar sites and Pazyryk this region’s population history, sample sizes are often small sites independently. All Tagar and Pazyryk samples date to and usually only contain information about the maternal an- the Late Bronze Age or Early Iron Age (based on artifacts cestry (mtDNA) of the individuals. In addition, sampling for found with the skeletal remains or carbon-14 dating of ob- ancient DNA is destructive. GMM methods allow research- jects found in the graves), and derive from locations in ers to study population history in a non-destructive context. southern Siberia (Alekseev et al., 2002; van Geel et al., Therefore, sample size is increased and a broader geograph- 2004). Pazyryk remains were not sampled from other loca- ic area can be included for comparison. GMM can also be tions that have associated remains, such as the Mongolian used to corroborate existing ancient DNA studies. Altai (Jordana et al., 2009). All analyses in this study include both Tagar and Pazyryk In order to assess population history and structure, we in- samples to understand how they are related to each other. clude both modern crania from Siberia and Central Asia That is, we do not analyze each group separately against our (18th–20th centuries) and pooled samples of ancient crania comparative dataset. If their close relationship persists when from Siberia, Mongolia, and China. We assume the prove- compared to geographically and temporally diverse samples, nance of our modern samples to derive from the last few then we can infer a close population history for these two centuries. Most of our ‘modern’ samples have documented groups. If either group appears more similar to other samples provenience. However, several samples from the Musée de used in the analysis, then we can infer something about be- l’Homme in Paris contain little or no provenance. The only tween group population structure and possibly external and information available to us was the geographic location from independent sources of gene flow. We suggest the Iron Age which the skulls were collected. Tagar and Pazyryk peoples will be more closely related to For instance, the linguist and ethnographer N.N. Pantusov, each other than they are to other peoples in the region based who excavated Kurgan remains dating from the Bronze on craniofacial variation. As phenotypic variation is akin to Age to modern times in Kazakhstan, also excavated our genetic distance, we should expect to see strong similarity sample labeled ‘Kazakhs’ during the late 1800s and early between these two ancient Siberian groups. 1900s, now curated at the Museé de l’Homme. These Ka- Our results show differential patterns of sex-biased migra- zakh crania were collected from a region historically known tion amongst Iron Age nomadic peoples of southern Siberia. as Semirechye, now known as Zhetysu in Almaty Province. We show that different patterns emerge between male and However, records indicate that the Semirechye Oblast, a female Pazyryk peoples. Male Pazyryk appear more closely former administrative locale of the Russian Empire, also in- related to Iron Age Tuvans and female Pazyryk are more cluded lands now part of northern Kyrgyzstan and adjacent isolated and diverse. Both male and female Pazyryk show provinces of Kazakhstan. Modern Kazakh peoples share similarity to the Kazakh sample. Iron Age peoples of the similar biological features with modern ethnic Kyrgyz. In Tuva region are not closely related to modern Tuvans as a our analysis, the Kazakh and Kyrgyz crania appear very result of large migrations of Iron Age Mongolians known as different. Therefore, we have, as a caveat, tentatively labeled Xiongnu. We show that Tagar males are more isolated from the Kazakh sample ‘modern,’ though they could most cer- other ancient southern Siberian groups and may represent tainly date to the medieval period (1100–1500 AD) or per- more recent migrants during the Iron Age, perhaps influ- haps even earlier. enced by groups residing in present day Central Asia, such We also use one sample, the Kalmyks, as representing the as Kazakhstan or Uzbekistan. Tagar females seem to share a northern Caucasus and Lower Volga region. The reason we population history with the Yakut of northeastern Siberia, include Kalmyks in the analysis is their recent western Mon- who speak a branch of the Turkic language. golian origin: they began migrating through the steppes of present days Kazakhstan and Central Asia at the end of 16th Materials and Methods century AD and settled in the northern Caucasus steppes and Volga basin in the 17th century AD (Nasidze et al., 2005). Phenotypic variability of crania was assessed using a suite All of our ‘ancient’ samples have sufficient provenance of 26 homologous landmarks that are common amongst cra- (Keyser-Tracqui et al., 2003; Tumen, 2006; Chikisheva, niometric studies and are used to assess overall shape and 2008; Zubova, 2008, 2013). Some of the samples used in the size differences among individuals (Table 1). This suite of present study have been used in previous studies of ancient landmarks includes traits associated with the upper face, population history and structure (Pozdnyakov, 2004; orbital areas, cranial vault, and occipital region. Cranial co- Moiseyev, 2006; Tumen, 2006; Chikisheva, 2008); however, 140 R.W. SCHMIDT AND A.A. EVTEEV Anthropological Science

Table 1. Homologous landmarks of the skull used for analyses Landmark Description* Nasion Intersection of the nasofrontal suture with the midsagittal plane. Ectoconchion Known as ‘EC’ in Bass (1995). The point where the orbital length line, parallel to the upper border, meets the outer rim. Zygoorbitale Point of articulation between the zygomaxillary suture and the lateral and inferior margin of the orbit. Zygomaxillare Most inferior point of the zygomaticomaxillary suture. Prosthion The most anterior point in the midline on the upper alveolar process. Bregma The intersection of the coronal and sagittal sutures, in the midline. Nariale (L) Bottom of nasal aperture. Measured at the most inferior point of the anterior nasal aperture. Nariale (R) Infranasion (left) Taken at the most superior point along the nasomaxillary suture. Infranasion (right) Simotic left Simotic measurements taken at the simotic chord, the minimum horizontal breadth of the two nasal bones, taken at the subtense. Simotic middle Simotic right Alare (L) Instrumentally determined most lateral point on the nasal aperture taken perpendicular to the nasal height. Alare (R) Frontomalare temporale (L) The most laterally positioned point on the frontomalar suture. Frontomalare posterior (L) The most posterior point on the frontomalar suture that is not in the temporal fossa. Frontomalare temporale (R) Frontomalare posterior (R) Glabella The most forward projecting point in the midline of the forehead at the level of the supra-orbital ridges and above the nasofrontal suture. Euryon (L) Point determined instrumentally at the most widely separated points on the two sides of the skull. Euryon (R) Basion The midpoint of the anterior margin of the foramen magnum most distant from the bregma. Opisthocranion The most posterior point on the skull not on the external occipital protuberance. Lambda The intersection of the sagittal and lambdoidal sutures in the midline. Inion The point at the base of the external occipital protuberance. * Descriptions from Bass (1995); Martin and Saller (1957); Brace and Hunt (1990); von Cramon-Taubadel (2009b).

Table 2. Bronze and Iron Age samples used in this study Sample name Males Females Location Site name Period Institution Analysis† Ancient Siberian Iron Siberia 16 14 Western Siberia Bystrovka-2 8th–6th centuries RASN CVA1, CVA2 BC Iron Tuva 8 11 Tuva Republic Arzhan-2; Dogehe-Baary II ~7th century BC RASN CVA1, CVA2 Medieval period 21 20 Western/Southern Tashara-Karier-2; Zarechno- ~1100–1500 AD RASN CVA1, CVA2 Siberia Ubinsky; Krjuchnoe-VI; Sanatornyj-1; Toropovo Pazyryk 25 20 Altai Gorny Altai, pooled series 5th–3rd centuries RASN CVA1, CVA2, BC CVA3 Tagar 7 10 Altai Minusinsk Basin, pooled series 7th–3rd centuries RASN CVA1, CVA2, BC CVA3 Ancient Mongolian Xiongnu 18 11 Mongolia Pooled Seies, mainly Central and 3rd century NUM CVA3 Western Mongolia BC–1st century AD Ancient China Xinjiang Bronze 11 9 China Tianshanbeilu, Hami Region ~3rd century BC JIDA CVA3 Totals 106 95 † Indicates specific canonical variates analysis in which the sample is included. See text for further details. JIDA, Research Center for Chinese Frontier Archaeology, Jilin University, Changchun, China; NUM, National University Mongolia, Ulaanbaatar, Mongolia; RASN, Institute of Archaeology and Ethnography, Russian Academy of Sciences, Siberian Branch, Novosibirsk, Russian Federation. Vol. 122, 2014 ANCIENT NOMADS OF SOUTHERN SIBERIA 141

Figure 2. Photographs of representative skulls for Pazyryk (A), Tagar (B) and Iron Age Tuva (C) peoples. All skulls are male. Pazyryk and Iron Age Tuva skull photos taken from Chikisheva (2008) and reprinted with permission from Elsevier.

Table 3. Modern samples used in this study Sample name Males Females Location Period Institution Analysis† Modern Siberia/Caucasus Buryat 10 11 Eastern Siberia Modern LMSU CVA2 Evenks 7 10 Eastern Siberia Modern LMSU CVA2 Kalmyk 17 17 North of Caspian Sea Modern LMSU CVA2 Orochi 8 9 East Siberia Modern LMSU CVA2 Tuva 19 19 Southern Siberia Modern LMSU CVA2, CVA3 Yakut 14 18 Northeastern Siberia CVA3 Ulchi 5 11 Eastern Siberia Modern LMSU CVA2 Modern Central Asian Kazakh 5 7 Central Asia Modern* MUSE CVA3 Kyrgyz 14 13 Central Asia Modern LMSU CVA3 Turkmen 13 4 Central Asia Modern LMSU, MUSE CVA3 Uighur 6 7 Western China Modern LMSU CVA3 Uzbek 9 7 Central Asia Modern LMSU, MUSE CVA3 Totals: 127 133 * Sample does not contain temporal provenience. See text for further details. † Indicates specific canonical variates analysis in which the sample is included. See text for further details. LMSU, Lomonosov Moscow State University, Moscow, Russian Federation; MUSE, Musée d l’Homme, Paris, France. our methodology (GMM coupled with multivariate analyses) (Xiongnu, pooled series from several sites in Central and has not been used previously. In addition, the sample com- Western Mongolia) that dates to the Iron Age (3rd century parisons we make have not been previously analyzed, with BC–2nd century AD) (Keyser-Tracqui et al., 2003; Schmidt, the exception of Chikisheva (2008), who included a more 2012). Lastly, we include a Bronze Age sample (Tianshan- diverse dataset with different aims (specifically uncovering beilu, Hami City) from the eastern Xinjiang Province (4th– the population history of the Tuvan peoples), and Moiseyev 3rd centuries BC) (Schmidt, 2012). We include this ancient (2006) who analyzed non-metric traits of the skull. sample as previous studies indicated some biological affinity The medieval Siberian sample (1100–1500 AD) contains among ancient Siberians and Bronze Age Xinjiang peoples pooled remains from several sites located in western and of the Tian Shan mountain range. This sample also included southern Siberia (Pozdnyakov, 2004). The other Early Iron distinct pottery closely linked to types seen in southern Sibe- Age Siberian sample (5th–3rd centuries BC) comes from ria and Western Mongolia (Liu and Cheng, 2012). one site (Bystrovka-2) in southern Siberia (Moiseyev, 2006). The raw data were analyzed using the software program The Iron Age Tuva sample (7th–3rd centuries BC) is pooled MorphoJ (Klingenberg, 2011). Landmark configurations from two sites (Arzhan-2 and Dogehe-Baary II) (Chikisheva, were processed by means of GMM (Klingenberg, 2010). 2008). In addition, we include a Turkic-Mongolian sample Original configurations were superimposed according to the 142 R.W. SCHMIDT AND A.A. EVTEEV Anthropological Science generalized Procrustes analysis (GPA) procedure in Mor- closely related to the Bystrovka-2 Iron Age females phoJ using the total covariance matrix. Sample variance was (Figure 3B). In both males and females, the pooled medieval assessed using canonical variates analysis (CVA) and Ma- series is separated along CV2. halanobis distances generated from the total covariance The Tagar and Pazyryk males and females were then com- matrix. In this study we do not attempt to interpret morpho- pared to modern Siberian crania (CVA2), including groups logical shape changes between samples, rather our goal is from northeastern Siberia (Evenks, Orochi, Ulchi) and the overall classification. Caspian Sea region (Kalmyks); and ancient Siberian crania, Three separate CVA analyses (CVA1, CVA2, CVA3) were mainly from southern and western Siberia (pooled medieval performed to assess ancient Siberian classification (for sam- sample, Iron Age Bystrovka-2, pooled Iron Age Tuva). The ples included in each analysis, see Table 1 and Table 2). In plot of the first two canonical variates accounts for 56.2% CVA1, we chose only representative samples from southern of the total variance for males, and 53.6% for females and western Siberia that date from the Iron Age to the medi- (Figure 4). The ellipses drawn around the population mean eval period. This was done in order to assess male and fe- are 75% frequency ellipses. The CVA plots for both males male distances among ancient Siberian groups. Differential (Figure 4A) and females (Figure 4B) indicate a general sep- patterns of sex-biased distances may indicate whether the aration of eastern Siberian groups and the Kalmyk sample local autochthonous component in southern Siberia was due compared to southern Siberian and ancient Siberian groups. to female or male groups. In CVA2, we chose both ancient However, female crania appear more diverse. For example, and modern Siberian groups. This analysis was performed in although there is still a general separation among ancient order to assess the apparent differences of the Iron Age and modern groups along CV1, the modern Tuvan females groups to modern northeastern Siberian groups. That is, we cluster tightly with the Kalmyk females. This is not the case included these groups intentionally to show the significant for modern male Tuvans, who appear in an intermediate po- differences among southern and eastern Siberian groups. sition; perhaps reflecting a mixed ancestry related to greater Based on previous research, there have been suggestions of gene flow from ancient groups compared with modern Tu- the influence of a Turkic-speaking people in southern Sibe- van females. In this analysis, the Tagar and Pazyryk male ria. Therefore, in CVA3, we included both male and female samples are more closely related due to the introduction of Pazyryk and Tagar samples and compared them with mod- more highly differentiated groups. However, the females for ern and ancient Turkic groups located in southern Siberia, both groups do not show a similar pattern. Although female Central Asia, and Western China. Tagar and Pazyryk are fairly close in Figure 4B, they do not overlap. The female Iron Age Tuvans are now more closely Results related to both female Tagar and female Pazyryk. Also, the Tagar males have become less isolated in this analysis. Male and female Tagar and Pazyryk craniofacial diversity It has been suggested that Central Asian Turkic (or Turk- was initially examined against several other contemporane- ish speaking) groups have played a role in shaping the genet- ous samples from the Iron Age in southern Siberia (CVA1), ic diversity seen today in southern Siberia. Therefore, the in addition to a pooled sample from the Siberian medieval Tagar and Pazyryk were analyzed against samples collected period (~1100–1500 AD). The CVA results show differential from Central Asia (Kazakh, Kyrgyz, Uighur, Uzbek, and patterns for Iron Age males and females. The plot of the first Turkmen), Siberia (modern Tuvans, a Turkic ethnic group), two canonical variates accounts for 86.4% of the total vari- a Mongol-Turkic Iron Age group from Mongolia (Xiongnu), ance for males (Figure 3A) and 71.7% for females. It is clear and a Bronze Age sample from Xinjiang (Xinjiang Bronze) from this plot (Figure 3A) that along CV1 Pazyryk males (CVA3). For male crania, the first two canonical variates cluster closer to Iron Age Tuvans, while the Tagar sample account for 48.8% of the variance (Figure 5A), while for and Iron Age males from the Bystrovka-2 site are more females, the first two canonical variates account for 53.8% isolated. On the other hand, female Pazyryk and Iron Age of the variance (Figure 5B). Similar to Figure 4, the ellipses Tuvans are separate along CV1, while the female Tagar are are 75% frequency ellipses. In Figure 5A, the Xinjiang

Figure 3. CVA plot for ancient male and female Southern/Western Siberians (CVA1). Closed circles are individual crania. (A) Male crania (n = 77; CV1 = 71.8%, CV2 = 14.6%). (B) Female crania (n = 75; CV1 = 47.9%, CV2 = 23.8%). Vol. 122, 2014 ANCIENT NOMADS OF SOUTHERN SIBERIA 143

Figure 4. CVA plot of the first two canonical variates of ancient and modern Siberian samples (CVA2). (A) Male crania (n = 142; CV1 = 40.2%, CV2 = 16.0%). (B) Female crania (n = 152; CV1 = 42.5%, CV2 = 11.1%). Closed circles are individual crania, ellipses are drawn as equal frequency ellipses (75%) around group mean.

Figure 5. CVA plot of the first two canonical variates for Pazyryk, Tagar and modern ethnic Turkic or ancient Turkic-speaking groups (CVA3). (A) Male crania (n = 140; CV1 = 31.9%, CV2 = 16.9%). (B) Female crania (n = 125; CV1 = 36.9%, CV2 = 16.9%). Closed circles are individual crania, ellipses are drawn as equal frequency ellipses (75%) around group mean. males are clear outliers along CV1. The Tagar males are also Table 4. Tagar and Pazyryk Mahalanobis distances for male mod- slight outliers, clustering loosely with the Pazyryk, modern ern Siberian/Caucasus cranial comparison Uzbeks, and the Kazakhstan sample. Interestingly, the fe- Buryat Evenks Kalmyk Orochi Pazyryk Tagar Tuva male CV plot shows a tight cluster of female Pazyryk, Tagar, Evenks 8.36 and Yakut to the exclusion of the other samples, which is not Kalmyk 6.38 8.56 shown among male crania (Figure 5B). Mahalanobis dis- Orochi 7.55 7.31 7.53 tances among modern Siberian and Central Asian groups Pazyryk 9.50 11.72 9.08 9.82 indicate more similarity of the Tagar and Pazyryk peoples to Tagar 10.16 12.30 10.53 10.66 6.55 some Central Asian samples as opposed to modern Siberian Tuva 7.42 11.05 9.12 9.89 7.47 8.44 samples (Table 4, Table 5, Table 6, Table 7). Although the Ulchi 9.29 7.63 9.83 7.66 11.35 12.41 10.81 result of NE Siberian groups separating from southern Sibe- rian groups is not surprising given the history of the region, the relative closeness of Kazakh male and female crania Discussion (that could date to as early as the medieval period) and male Xiongnu sample to Pazyryk and Tagar could indicate ancient Southern Siberian groups, both modern and ancient, have admixture or gene flow among these groups. been investigated extensively, owing to a rich and diverse cultural hybrid zone that has seen significant contact among peoples originating from both Western and Eastern Eurasia 144 R.W. SCHMIDT AND A.A. EVTEEV Anthropological Science

Table 5. Tagar and Pazyryk Mahalanobis distances for male Turkic cranial comparison Xinjiang Kazakh Kyrgyz Pazyryk Tagar Turkmen Tuva Uighur Uzbek Xiongnu Bronze Kyrgyz 11.11 Pazyryk 6.55 8.78 Tagar 10.39 11.93 8.48 Turkmen 9.09 7.34 6.59 9.34 Tuva 8.58 8.15 7.50 9.74 8.55 Uighur 9.17 9.47 7.85 11.06 6.45 8.64 Uzbek 7.32 9.08 6.63 7.83 6.77 7.66 7.87 Xinjiang Bronze 10.51 13.91 9.25 9.43 12.57 11.49 12.72 10.41 Xiongnu 7.73 9.13 6.23 9.91 8.84 7.02 9.49 7.73 9.59 Yakut 7.44 9.62 6.99 10.15 8.80 8.12 10.62 7.35 10.59 6.81

Table 6. Tagar and Pazyryk Mahalanobis distances for female peoples of Siberia and Central Asia (Figure 5), while main- modern Siberian/Caucasus comparison taining a connection to some peoples from the Iron Ages of Buryat Evenks Kalmyk Orochi Pazyryk Tagar Tuva southern and western Siberia (Figure 3, Figure 4, Figure 5). These findings are similar to previous research done in an- Evenks 7.62 cient DNA studies and physical anthropological studies Kalmyk 6.81 7.35 (Chikisheva, 2000a; Moiseyev, 2006; Chikisheva et al., Orochi 8.32 7.30 8.09 2007; Pilipenko et al., 2010). Pazyryk 10.25 10.68 7.37 10.48 It is well known that the modern genetic diversity seen in Tagar 11.04 12.15 8.99 12.49 6.57 the region of southern Siberia today stems from extensive Tuva 7.87 8.20 6.43 8.62 8.16 10.54 contacts among diverse peoples (Comas et al., 1998, 2004). Ulchi 8.06 7.22 7.92 7.01 10.66 12.84 7.63 Mitochondrial DNA and Y chromosome studies have been conducted on a range of living peoples in Northern Asia, including Tuvinans, Buryats, Khakassians, Sojots, Todjins, since as early as the Upper Paleolithic. Importantly, this Tofalars, Kalmyks, Kazakhs, Kizhi, Mongols, Evenks, and region has been hypothesized to be the origin of modern-day Yakuts, among others (Kolman et al., 1996; Wells et al., Native American peoples, while also maintaining high levels 2001; Zerjal et al., 2002; Derenko et al., 2003, 2006, 2007; of genetic and cultural diversity (Quintana-Murci et al., Pakendorf et al., 2003, 2006; Nasidze et al., 2005; 2004; Dulik et al., 2012; Raghavan et al., 2013). This study Starikovskaya et al., 2005; Phillips-Krawczak et al., 2006; has attempted to reconcile the population history of southern Gokcumen et al., 2008). These groups range in linguistic Siberia by examining a small temporal slice during the Iron diversity as well, covering Turkic, Mongolic, and Tungusic Age. Notably, two important cultural groups with questiona- language families. The majority of these studies include ble origins were investigated craniometrically to elucidate analyses of haplogroup reconstruction for the uniparentally questions owing to divergent morphological appearances inherited mtDNA and Y chromosome. The general consen- and mtDNA haplotype composition (Voevoda et al., 1998; sus among these studies is that the mountain belt zone of Chikisheva et al., 2007). These cultural groups, the Tagar southern Siberia is where populations began to expand into and Pazyryk, are important to understanding the broader Eastern and Northern Europe following the Last Glacial history of the region. Our results have shown these two Maximum. From the Mesolithic, the region of southern groups to be outliers when compared to many modern living Siberia has witnessed extensive migrations, most notably

Table 7. Tagar and Pazyryk Mahalanobis distances for female Turkic cranial comparison Xinjiang Kazakh Kyrgyz Pazyryk Tagar Turkmen Tuva Uighur Uzbek Xiongnu Bronze Kyrgyz 11.53 Pazyryk 8.44 9.80 Tagar 8.99 10.62 5.95 Turkmen 11.14 8.97 11.81 12.35 Tuva 9.56 8.72 8.20 10.79 10.38 Uighur 10.07 8.70 7.75 8.85 9.53 9.37 Uzbek 9.90 10.83 8.61 8.98 11.83 11.52 8.59 Xinjiang Bronze 12.14 16.74 11.08 10.15 17.02 14.47 14.16 13.72 Xiongnu 10.09 13.53 8.84 9.86 13.21 10.62 11.81 13.32 10.76 Yakut 7.59 13.13 8.11 9.02 12.67 11.21 10.41 9.62 12.11 9.26 Vol. 122, 2014 ANCIENT NOMADS OF SOUTHERN SIBERIA 145 during the Bronze and Iron Ages. (2004), who found that lineages in Kazakhstan before the Paleogenetic studies of the southern Siberian region dur- Iron Age all belonged to Western Eurasian lineages. During ing the Bronze and Iron Ages have included mtDNA and Y the Late Iron Age, the influx of Turko-Mongolian Xiongnu chromosome analyses of Pazyryk, Xiongnu, Scythian, increased the Eastern Eurasian lineages present in Kazakh- Tagar, and Kazakhstan peoples (Clisson et al., 2002; Keyser- stan (Lalueza-Fox et al., 2004). Therefore, the Tagar male Tracqui et al., 2003; Lalueza-Fox et al., 2004; Ricaut et al., outlier position in Figure 3 may be the result of incoming 2004a, b; Chikisheva et al., 2007; Keyser et al., 2009; East Eurasian peoples during the Late Iron Age. Pilipenko et al., 2010; Gonzalez-Ruiz et al., 2012). The gen- More generally, our results show some population differ- eral consensus among these studies is that the region of ences between the Pazyryk and Tagar peoples. As seen in southern Siberia and Central Asia is quite diverse. However, Figure 3, the two Iron Age groups do not overlap for the first studies have shown that Pazyryk and Tagar peoples pos- two canonical variates when compared with a small number sessed haplogroups that are rare or absent in southern Siberia of contemporaneous groups. In fact, the Tagar males seem to today, such as mtDNA haplogroups U and U5a1 (Pilipenko be greater outliers than the Pazyryk while we can see the et al., 2010). Interestingly, haplogroup U is present in high opposite situation for females of both groups. The Pazyryk frequencies in ancient hunter-gatherers of Europe (Bramanti in Figure 3 overlap with, and are close to the Iron Age Tu- et al., 2009; Malmstrom et al., 2009) and was also found in a vans, and to a lesser degree to other Siberian Iron Age sam- 24000 year old Upper Paleolithic boy from the Lake Baikal ples. Therefore, the connection between the Pazyryk peoples region (Mal’ta) of south-central Siberia (Raghavan et al., in Siberia may be stronger than for Tagar peoples. 2013). Haplogroup U5a1 has also been observed in eastern The situation for female groups looks to be otherwise: the Kazakhstan from the Bronze Age (Lalueza-Fox et al., 2004). Pazyryk females seem to be outliers while Tagar females are Therefore, the connection between pre-agricultural Europe- similar to the Iron Age Siberia sample from Bystrovka-2. an peoples and the Pazyryk peoples may have been estab- Taken together, these results point to a high probability of lished from as early as the pre-Neolithic. sex-biased admixture in both groups where migrating Tagar Our craniofacial study did not include peoples of modern- males assimilated some previous Iron Age population while day Europe with high frequencies of haplogroup U or other Pazyryk males are more similar to neighboring Iron Age typical Western Eurasian haplogroups such as R (Quintana- Tuvan peoples who may have had other sources of marital Murci et al., 2004). We did, however, observe stark differ- partners. Chikisheva (2008) showed that the nomads of Iron ences between the Iron Age Siberians and modern Siberians Age Tuva formed a subcluster with the pooled Pazyryk sam- that contain higher frequencies of common Eastern Eurasian ple, but not the Tagar peoples. This result is clearly seen in haplogroups, such as M (Derenko et al., 2012; Gonzalez- our CVA shown in Figure 3, but only for the male sample Ruiz et al., 2012). Our results would therefore suggest the while the females obviously differ from the Iron Age Tuvan Pazyryk and Tagar peoples either had greater contact with group, at least on CV1. It should be noted that the Iron Age West Eurasian peoples, or their ancestors originated or had Tuvan sample is the closest group for both males and fe- close connections with the hunter-gatherers of Paleolithic males of Tagar. However, this result should not be overem- Europe. This observation is reinforced through ancient DNA phasized as the comparison here is done among very few analysis. Keyser et al. (2009) analyzed mtDNA and Y chro- groups of generally common origin and cranial morphology. mosome haplotypes of Middle and Late Bronze Age, and After introducing several northeastern Siberian groups Iron Age southern Siberian samples, including samples from into the analysis (Figure 4), the Tagar and Pazyryk males do the Tagar culture. Their results revealed that all of their sam- not differ greatly and strongly overlap, while females of both ples contained the Y haplogroup R1a1, which is widely dis- groups are still fairly distinguishable and do not overlap, in- tributed on the Eurasian continent (Karafet et al., 2008). This dicating greater heterogeneity among ancient southern Sibe- haplogroup most likely reflects the expansion of peoples af- rian females. Overall, females of the Iron Age and medieval ter the Last Glacial Maximum (~20–12 kya) and therefore southern Siberian groups look more diverse compared to the Bronze and Iron Age peoples of Siberia were part of a males of the same groups even against the background of continuation of West Eurasian peoples that may have such morphologically different groups such as Northeast reached as far as Lake Baikal during the Upper Paleolithic Asians (Figure 4B). (Raghavan et al., 2013). However, when we compared the Pazyryk and Tagar sam- While population genetic studies point mostly to Western ples to crania collected from Central Asian groups (Kazakhs, (European) direction of relationships of Tagar and Pazyryk Turkmen, Uzbeks, Uighurs) with the addition of ancient peoples, both archaeological and craniological data suggest Xiongnu, who originated in Mongolia and date to the Iron fairly strong influence of Central or even East Asian popula- Age, and a Bronze Age sample from Xinjiang (Figure 5), we tions on the Iron Age tribes of southern Siberia (Chikisheva, found the Pazyryk males to be more closely related to Cen- 2000a, 2008). It is known from ancient DNA studies (Keyser tral Asian groups than Tagar males, who show an outlier et al., 2009; Pilipenko et al., 2010) that western and southern position though fairly close to Pazyryk males (Figure 5A). Siberian Bronze Age samples harbored higher frequencies On average, both the Tagar and Pazyryk peoples are more of Western Eurasian than Eastern Eurasian mtDNA hap- closely related to Central Asian groups, namely the Uzbek logroups prior to the Iron Age. In the Keyser et al. (2009) sample and Pantusov’s collection from Kazakhstan (see Ma- study, the frequency of Western Eurasian mtDNA hap- terials and Methods section for details of this sample prove- logroups reached 90%, but lowered to 67% during the Iron nance) than modern northern Siberian groups. Interestingly, Age. This is similar to the results of Lalueza-Fox et al. when the ancient Xiongnu and the Bronze Age Xinjiang are 146 R.W. SCHMIDT AND A.A. EVTEEV Anthropological Science included in the analysis, neither the Tagar nor Pazyryk peo- lic of China, Philippe Mennecier at the Musée de l’Homme, ples overlap with individuals from the Xiongnu sample Paris, France, and Tumen Dashtseveg at the National Uni- (Figure 5). Thus, a direct genetic influx from the Xiongnu versity of Mongolia, Ulaabaatar. The first author would also population does not look very probable in either the Tagar or like to thank Mary Margaret-Murphy at the University of Pazyryk peoples. Montana for help with data formatting, Pauline Sebillard at Since the Pazyryk and Tagar samples are pooled from Jilin University for translating in Changchun, and Natasha different sites and different time periods, their affinity to the Kharlamova at the Russian Academy of Sciences for help in Central Asian Kazakh sample could stem from later incur- Moscow. We would also like to thank the three anonymous sions of Eastern Eurasian peoples toward the end of the Iron reviewers and the Editor’s comments in helping to greatly Age. These results have not been demonstrated previously, improve the final version of this manuscript. Research was though Chikisheva (2000a) did observe for both male and partially supported through a National Science Foundation female Pazyryk crania a close biological similarity to the Doctoral Dissertation Improvement Grant (BCS no. 1028773) ancient Saka and Wusun groups of eastern Kazakhstan and (R.W.S.). Xinjiang. Unfortunately, to our knowledge, we were unable to include ancient samples from Kazakhstan, however, we do show a similar pattern for our pooled Pazyryk sample, References which could indicate that our Kazakh sample dates from Alekseev A., Bokovenko N.A., Boltrik Y., Chugunov K.A., Cook much earlier than the last few centuries. Importantly, the G., Dergachev V.A., Kovaliukh N., Possnert G., Der Plicht J., Kazakh sample is the only one showing similarity to both Scott E.M., Sementsov A., Skripkin V., Vasiliev S., and Zaitseva sexes of (Tagar and Pazyryk) cultures. Pazyryk females, G. 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(1998) Trading genes along the Silk Road: mtDNA sequences and the origin of Central We would like to thank the museum curators and individ- Asian populations. American Journal of Human Genetics, 63: uals from the institutions where skeletal material is curated. 1824–1838. This includes Alisa Zubova at the Institute of Archaeology Comas D., Plaza S., Wells R.S., Yuldaseva N., Lao O., Calafell F., and Bertranpetit J. (2004) Admixture, migrations, and disper- and Ethnography, Russian Academy of Sciences, Siberian sals in Central Asia: evidence from maternal DNA lineages. Branch, Novosibirsk, Russian Federation, Zhu Hong and European Journal of Human Genetics, 12: 495–504. Wei Dong at the Research Center for Chinese Frontier Ar- Derenko M.V., Grzybowski T., Malyarchuk B.A., Dambueva I.K., chaeology of Jilin University, Changchun, People’s Repub- Denisova G.A., Czarny J., Dorzhu C.M., Kakpakov V.T., Vol. 122, 2014 ANCIENT NOMADS OF SOUTHERN SIBERIA 147

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