Prehistoric and Recent Populations of Chukotka: A Paleophenetic Analysis

Prehistoric and Recent Populations of Chukotka: A Paleophenetic Analysis

ARCHAEOLOGY, ETHNOLOGY & ANTHROPOLOGY OF EURASIA Archaeology Ethnology & Anthropology of Eurasia 40/3 (2012) 130–137 E-mail: [email protected]...

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ARCHAEOLOGY, ETHNOLOGY & ANTHROPOLOGY OF EURASIA Archaeology Ethnology & Anthropology of Eurasia 40/3 (2012) 130–137 E-mail: [email protected]

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ANTHROPOLOGY A.A. Movsessyan Moscow State University, Leninskie Gory 1, building 12, Moscow 119991, Russia E-mail: [email protected]

PREHISTORIC AND RECENT POPULATIONS OF CHUKOTKA: A PALEOPHENETIC ANALYSIS

Thirty nonmetric traits were studied in two cranial series from prehistoric coastal cemeteries on the Chukchi Peninsula (hereafter Chukotka) – Welen (58 crania) and Ekven (107 crania), representing the Old Bering Sea culture. Both series are close not only to modern Eskimos, Chukchi, and Aleuts but also to Tungus-speaking groups. This may be due both to ancient contacts between continental and coastal populations and to their common origin. Comparisons with the Neolithic groups of Baikal and with southern Mongoloids suggest that Eskoaleuts originated in Siberia, and that admixture between migrants from continental Siberia and those from more southern areas of the Paci¿c coast was one of the key factors in this process. Another factor was adaptation to the extreme Arctic environment. Keywords: Craniology, population history, nonmetric cranial traits, Chukchi Peninsula, Eskimos, Aleuts, Chukchi.

Introduction The purpose of this study is to assess the position of cranial series from two pre-Metal Age cemeteries on the Beringian coast of Chukotka – Welen and Ekven – among other series from Northern Asia, using nonmetric traits. These two groups, representing the Old Bering Sea culture, were ¿rst studied by M.G. Levin (1962), who, based on craniometric data, described them as quite Eskimo-like, concluding that the Eskimo trait combination was very ancient. G.F. Debetz (1975) reached the same conclusion in his study of Eskimo origins. The dental analysis revealed that both groups were similar to each other and to recent Eskimos, Chukchi, and Aleuts, and that both groups had retained ancient trait combinations of the dental structure found in certain Mesolithic and Neolithic populations (Zubov, 1969).

Results of craniometric comparisons between ancient and recent groups are often affected by diachronic variation, which distorts population relationships. These relationships can be more reliably reconstructed if the traits are unaffected by temporal trends. This appears to be the case with nonmetric characters of the cranium. In this study, they provide a basis for a comparison of prehistoric samples from Chukotka with modern Mongoloid groups. Nonmetric traits, known as “phenes” in Russian zoological literature, are used as quasi-genetic markers by zoologists in population studies. Principles of population genetics are thereby extended to cases where genetic analysis is dif¿cult or impossible (Timofeyev-Resovsky, Yablokov, 1973; Timofeyev-Resovsky et al., 1973). Indirect data suggests that nonmetric variations of the cranium result from normal developmental processes and are to some extent controlled by genetic factors

© 2012, 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.2012.11.015

A.A. Movsessyan / Archaeology, Ethnology and Anthropology of Eurasia 40/3 (2012) 130–137

(Lane, 1977; Cheverud, Buikstra, 1981; Sjøvold, 1984). Also, numerous studies have demonstrated that groups unrelated by origin differ in the frequencies of these traits, whereas the likelihood of related groups being similar is much higher. Therefore the application of genetic methods appears to be very useful in the analysis of population af¿nities between ancient groups. Materials and methods In our earlier publications we have proposed a list of nonmetric cranial traits and a methodology of their

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analysis (Movsessyan et al., 1975; Movsessyan, 2005). In this study, these traits were scored on 107 crania from Welen and 58 from Ekven. For comparative purposes, their frequencies in previously published prehistoric and recent Mongoloid series were used (Mamonova, Movsessyan, 1998; Movsessyan, 2005). Frequencies were calculated per individual, i.e. without regard to unilateral or bilateral occurrence. Statistical analysis was conducted using three methods: (1) Nei’s generalized distance (d) (Nei, 1972); (2) classification analysis based on the PHYLIP – Phylogeny Inference Package (Felsenstein, 1989); and (3) canonical variate analysis based on the CANOCLUS package by V.Y. Deryabin.

Table 1. Frequencies of nonmetric cranial traits in ancient populations of Chukotka and modern Eskimos, Chukchi, and Aleuts No.

Traits

Welen

Ekven

Eskimos

Coastal Chukchi

Reindeer Chukchi

Aleuts

1

Sutura frontalis

0.051

0.028

0.049

0.040

0.010

0.054

2

Foramen supraorbitale

0.579

0.443

0.637

0.733

0.777

0.642

3

Foramen frontale

0.129

0.141

0.196

0.160

0.167

0.071

4

Spina trochlearis

0.010

0.019

0.027

0.010

0.010

0.038

5

Foramen infraorbitale accessorium

0.120

0.129

0.151

0.177

0.196

0.250

6

Os zygomaticum bipartitus trace

0.021

0.010

0.029

0.040

0.028

0.167

7

Spina processus frontalis

0.069

0.205

0.315

0.266

0.171

0.163

8

Os Wormii suturae coronalis

0.010

0.018

0.010

0.010

0.010

0.010

9

Stenocrotaphia

0.043

0.108

0.243

0.134

0.152

0.085

10

Os epiptericum

0.108

0.166

0.045

0.067

0.057

0.100

11

Processus frontalis squamae temporalis

0.010

0.090

0.010

0.013

0.014

0.020

12

Os Wormii suturae squamosum

0.021

0.019

0.010

0.010

0.010

0.020

13

Os postsquamosum

0.103

0.151

0.127

0.120

0.180

0.089

14

Os asterion

0.034

0.122

0.029

0.053

0.029

0.054

15

Foramen parietale

0.327

0.383

0.513

0.400

0.416

0.446

16

Os Incae

0.017

0.047

0.010

0.040

0.014

0.019

17

Os triquetrum

0.030

0.038

0.045

0.053

0.014

0.058

18

Os apicis lambae

0.034

0.010

0.010

0.014

0.027

0.038

19

Os Wormii suturae lambdoideae

0.086

0.113

0.063

0.085

0.083

0.192

20

Os Wormii suturae occipito-mastoideum

0.010

0.010

0.049

0.160

0.039

0.073

21

Processus interparietalis

0.086

0.047

0.039

0.069

0.069

0.010

22

Tuberculum praecondylaris

0.050

0.047

0.050

0.046

0.059

0.022

23

Foramen tympanicum

0.175

0.184

0.167

0.236

0.222

0.178

24

Foramen pterygospinosum

0.034

0.037

0.078

0.093

0.062

0.072

25

Foramen pterygo-alare

26

Sutura palatina transversa (concave)

27

Torus palatinus

0.185

0.208

0.147

0.155

0.223

0.286

28

Torus mandibularis

0.346

0.220

0.075

0.250

0.050

0.238

29

Sulcus mylohyoideus

0.154

0.110

0.125

0.125

0.142

0.142

30

Foramen mentale accessorium

0.077

0.060

0.050

0.250

0.285

0.200

0

0

0.039

0.046

0.078

0.036

0.018

0.019

0.061

0.111

0.061

0.036

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Results and discussion The resemblance between Welen and Ekven, demonstrated by craniometric and dental analyses, is likewise revealed by the analysis of nonmetric cranial traits. The generalized distance between them is small (d = 0.030). Apparently, both groups represent local populations of the same ethnic group. Both groups are close to recent Eskimo, Chukchi, and Aleuts (Table 1). Importantly, this similarity primarily concerns traits distinguishing these groups from other Northern Asian Mongoloids. For instance, crania from Welen and Ekven, like those of modern Eskimos, Chukchi, and Aleuts and unlike those of continental Siberian Mongoloids, are characterized by a high frequency of infraorbital and frontal foramina and of mandibular tori (the latter trait is especially typical of Eskimos, Chukchi, and Aleuts). On the other hand, frequencies of trochlear spines, lambda bones, and lambdoid Wormians are rather low in all these groups. The similarity is apparent from generalized distances (d) (Table 2). On the other hand, both prehistoric populations of Chukotka resemble the Tungus-speaking people of the Amur – the Ulchi and Negidal. Physical traits suggestive of Tungus af¿nities have been described in modern reindeer and coastal Chukchi (Levin, 1958). The same af¿nities are

revealed by nonmetric cranial traits (Table 3). Generalized distances show the Eskimos to be especially close to coastal Chukchi. Both groups of Chukchi (reindeer and coastal) are rather similar to each other. The Aleuts take a separate position (see below). Chukchi and Eskimos, then, form a cluster separated by moderate distances not only from Aleuts but also from Negidal and Ulchi. This is even more apparent in a plot showing the position of modern Siberian populations on two canonical vectors (Fig. 1). The Tungus-speaking groups – Ulchi, Negidal, Orochi, and Evenki – form a distinct cluster and, together with the Eskimos, Chukchi, and Aleuts, score positively on the ¿rst canonical variate. The results of two independent statistical methods, therefore, are in agreement. They suggest that af¿nities between Chukotkan populations and those speaking Tungus languages are quite ancient (no later than the Old Bering Sea culture and possibly even earlier). This conclusion is supported by ancient cultural links between Chukotka and Eastern Siberia. S.I. Rudenko (1947) suggested that the Old Bering Sea culture might have had a southern origin. Speci¿cally, the curvilinear design on artifacts from ancient Chukotkan cemeteries is close to that typical of the Amur area. It was hypothesized that the Old Bering Sea culture resulted from a blend of paleoEskimo cultures with labrettes and the Ust-Belaya culture,

Table 2. Distances between ancient Chukotkans and modern Siberian groups

Table 3. Distances between Eskimos, Chukchi, and Aleuts and populations of inland Siberia

Groups

Welen

Ekven

Chukchi

Ancient Chukotkans, pooled

Groups

Eskimos

Coastal Reindeer

Aleuts

Eskimos

0.051

0.061

0.050

Coastal Chukchi

0.029

0.000

Coastal Chukchi

0.050

0.068

0.052

Reindeer Chukchi

0.035

0.022

0.000

Reindeer Chukchi

0.041

0.071

0.047

Aleuts

0.070

0.047

0.056

0.000

Aleuts

0.044

0.059

0.046

Negidal

0.055

0.051

0.056

0.062

Negidal

0.044

0.051

0.041

Ulchi

0.040

0.039

0.055

0.034

Ulchi

0.048

0.055

0.043

Orochi

0.070

0.050

0.069

0.039

Evenki

0.056

0.075

0.060

Evenki

0.065

0.059

0.073

0.039

Orochi

0.071

0.074

0.069

Beltyr

0.084

0.092

0.130

0.075

Selkup

0.075

0.071

0.066

Sagay

0.109

0.104

0.120

0.089

Beltyr

0.143

0.127

0.130

Shorians

0.080

0.088

0.113

0.058

Sagay

0.111

0.090

0.096

Koibal

0.078

0.115

0.140

0.075

Shorians

0.098

0.077

0.079

Kachin

0.077

0.073

0.082

0.033

Koibal

0.109

0.081

0.090

Telengit

0.092

0.085

0.099

0.052

Kachin

0.076

0.085

0.073

Byryat

0.072

0.041

0.071

0.044

Telengit

0.101

0.106

0.095

Tuvans

0.081

0.052

0.071

0.038

0.072

0.078

0.096

0.043

0.066

0.074

0.087

0.044

Buryat

0.099

0.115

0.100

Mansi

Tuvans

0.068

0.093

0.075

Khanty

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which lacked labrettes. By the end of the 1st millennium BC, the Ust-Belaya culture spread toward the northern and eastern coast of Chukotka. This culture displays both Eskimo features and those typical of inland Siberia (Dikov, 1974). I.I. Gokhman (1961), who studied a cranium from UstBelaya cemetery, concluded that it evidenced af¿nities with Eskimos, Chukchi, and Aleuts, on the one hand, and Tungus-speaking groups, on the other. V.P. Alekseyev (1967) described the Ust-Belaya cranium as “protoEskimoid,” i.e., showing a more generalized morphology compared to that of extant Eskimos. Therefore our result indicating biological affinities between ancient Chukotkans and the Tungus speakers may attest both to contacts and to a common origin. An important aspect of group differentiation is “genetic memory,” implying that extant populations related by origin, preserve various characteristics of the ancestral population. This feature of population systems was first discovered by A.S. Serebrovsky (1935) and was later experimentally demonstrated by Y.G. Rychkov and Y.P. Altukhov (Rychkov, 1969, 1973; Altukhov, Rychkov, 1970; Rychkov, Movsessyan, 1972; Altukhov, Pobedonostseva, 1978). Under this approach, a population system is regarded as a subdivided population having its own microevolutionary history and consisting of subpopulations related by origin. The studies cited above have convincingly demonstrated that such a system is genetically stable in both time and space. Ethnic differentiation, because of being linked to genetic differentiation, follows the same pattern. Therefore ethnic groups, too, may preserve “memories” of ancestral communities despite being separated from them by dozens of generations. By averaging gene (and phene) frequencies across extant populations and ethnic groups one may arrive at a reconstruction of the ancestral group, thus offering the possibility for comparing modern populations with ancient ones. Using this approach, we can assume that the characteristics of modern Siberian groups and those of ancient groups of Chukotka to some extent mirror those of the hypothetical ancestral group. If so, comparing modern averages is tantamount to comparing ancestral populations. Such a comparative analysis has revealed that the ancient inhabitants of Chukotka were virtually

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Fig. 1. Results of the canonical variate analysis of nonmetric cranial traits in modern Siberian populations.

equally close to the ancestors of modern Eskimos, Chukchi, and Aleuts, on the one hand, and those of the Tungus speakers, on the other (Table 4). This may attest to southwestern af¿nities of the ancient Chukotkans. However, populations of inland Siberia including the Tungus speakers are much closer to the Neolithic population of Baikal. In a dendrogram, ancient Chukotkans cluster with modern Eskimos, Chukchi, and Aleuts (or, under the approach outlined above, with their common ancestors), whereas continental Siberians join the Baikal Neolithic groups (Fig. 2). The continuity between ancient and modern populations is especially apparent in diagrams showing the combinations of frequencies of the ¿fteen most polymorphic traits in separate groups and in reconstructed ancestral groups (Fig. 3). In sum, our results suggest that modern Eskimos, Chukchi, and Aleuts are not only similar to each other in terms of nonmetric cranial traits, but retain the “memory” of their prehistoric ancestors – people of the Old Bering Sea culture. This culture, in turn, was a stage in the evolution of the paleo-Eskimo sea hunters, which originated in the 2nd millennium BC on the Beringian coast of Chukotka as an adaptation to a speci¿c environment. The paleo-Eskimos are believed to be descendents of Late Paleolithic proto-Eskoaleuts, who in the Late Pleistocene and early Holocene spread across the

Table 4. Distances between ancient groups and pooled modern groups of Siberia Groups

Eskimos, Aleuts, Chukchi

Tungus-speaking groups

Western Siberians

Turks and Mongols

Welen

0.035

0.037

0.065

0.061

Ekven

0.048

0.046

0.053

0.074

Ancient Chukotkan, pooled

0.034

0.036

0.052

0.062

Baikal Neolithic

0.038

0.028

0.024

0.025

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Fig. 2. Dendrogram based on pairwise differences between modern and ancient Siberian populations.

Fig. 3. Graphs showing patterns of frequencies of the ¿fteen most polymorphic traits in separate groups and in reconstructed ancestral groups. Frequencies of traits plotted along the radii, clockwise: sutura frontalis, foramen frontale, spina processus frontalis, stenocrotaphia, os epiptericum, os postsquamosum, os asterion, os apicis lambdae, os Wormii suturae lambdoideae, os Wormii suturae occipito-mastoideum, processus interparietalis, tuberculum praecondylaris, foramen tympanicum, torus palatinus, sulcus mylohyoideus. The center of the circle corresponds to zero, the circle, to 0.25.

Beringian landbridge from Asia to northwestern America (Dikov, 1974). The genetic sources of the Old Bering Sea people are obscure. We can try, following A.M. Zolotarev (1937), to trace these sources in continental Siberia. Regrettably,

ancient skeletal materials from territories immediately adjoining Chukotka are quite fragmentary. It appears warranted, however, to compare ancient Chukotkans with people of the Baikal Neolithic, dated to 7th– 2nd millennia BC. The results demonstrate that in terms of nonmetric trait frequencies, the “paleo-Eskimos” differ from the Neolithic people of the Baikal no more than from modern Eskimos, Chukchi, and Aleuts, being closest to the earliest group – Kitoy (7th–6th millennia BC) (Table 5). Graphs showing patterns of trait distribution in ancient populations of Chukotka and Baikal, demonstrate Serovo and Glazkovo groups as resembling one another while differing from the Kitoy group, which is to some extent close to the reconstructed ancestral Chukotkan group (Fig. 4). Results of the canonical variate analysis of modern and ancient populations of Siberia fully agree with the above results and supplement them (Fig. 5). Modern Eskimos, Chukchi, Aleuts, and Tungus-speakers as well as Welen, Ekven, and Kitoy show positive scores on the second canonical variate. Modern Turks, Mongols, and Uralians, on the other hand, join Serovo and Glazkovo (they all score negatively on this CV). Notably, Aleuts differ from Eskimos and Chukchi and resemble inland North and Central Asians. The uniqueness of Aleuts has already been noted. Being culturally and linguistically close to Eskimos, Aleuts are anthropometrically distinct from them (Tokareva, 1937; Hrdliþka, 1944; Debetz, 1951; Rychkov, Sheremetyeva, 1972; Sheremetyeva, Rychkov, 1978; Alekseyev, 1981). Whereas G.F. Debetz believed that the Aleuts were related to Eskimos and Chukchi, N.N. Cheboksarov (1947) linked them to Buryats, Tuvans, and Mongols. Results of nonmetric cranial analysis indicate the same. Speci¿cally, canonical variate analysis shows the Aleuts to be much closer to Tuvans and Evenki than to Eskimos and Chukchi (Fig. 1, 5). If the ancestors of ancient Chukotkans lived in Central Siberia, the biological peculiarities of Aleuts can be explained by isolation, which contributed to the conservation of traits of a population that was ancestral to proto-Eskimos and Aleuts, and was close to ancient Central Siberians. According to T.Y. Tokareva, who discussed craniometric af¿nities between Aleuts, Evenki, and the Neolithic Baikalians, Aleut ancestors

Table 5. Distances between ancient Chukotkans and the Baikal Neolithic groups Kitoy

Serovo

Glazkovo

Baikal Neolithic, pooled

Welen

0.035

0.047

0.051

0.037

Ekven

0.041

0.05

0.038

0.036

Ancient Chukotkans, pooled

0.032

0.043

0.039

0.031

Groups

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Fig. 5. Results of the canonical variate analysis of nonmetric cranial traits in modern and ancient Siberian populations.

Fig. 4. Graphs showing patterns of trait frequencies in ancient populations of Chukotka and Baikal. See Fig. 3 for explanations.

were related to Neolithic Baikalians. She believed that sometime in the past those people spread across a vast territory that included the Bering Sea Basin and possibly northwestern America (Tokareva, 1937). In support of this view, V.P. Alekseyev wrote that the morphophysiological peculiarities of modern Aleuts could be due to the conservation of proto-Mongoloid traits (Alekseyev, Trubnikova, 1984: 75). Indeed, in terms of cranial nonmetrics, Aleuts are rather close to the pooled Neolithic group from Baikal (d = 0,036). If ancient populations are included in the canonical variate analysis, people of Welen and Ekven shift toward modern Eskimos, Chukchi, and Aleuts (Fig. 6), in agreement with the results shown in the dendrogram (Fig. 2). The Neolithic Baikalians, on the other hand, shift toward the center of the continental Siberian cluster. Thanks to the inclusion of the supposed ancestral Chukotkans, Aleuts cluster with Eskimos, and Chukchi, showing positive scores of the second canonical variate and negative scores of the ¿rst. They are equally removed from the Neolithic Baikal group and from modern Turks and Mongols. It can be concluded that the average characteristics of ancient populations indeed reflect those of ancestral groups: proto-Eskoaleut and protoContinental Siberian. Apparently, proto-Eskoaleuts, like the Neolithic Baikal populations, originated from the same proto-Mongoloid Paleolithic population of Siberia which, in the course of dispersal and ¿ssion, had given rise to local Neolithic populations.

Fig. 6. Results of the canonical variate analysis of nonmetric cranial traits in modern and ancient Siberian populations (pooled ancient groups added).

Our results are supported by archaeological data, suggesting that the Neolithic culture of the Angara and Baikal began to expand in the late 4th millennium BC. The Lena–Aldan Neolithic culture spread via the upper Lena, down the Angara, and further northwest and north, to Khatanga and the lower Lena as well as eastwards along the Okhotsk Sea coast toward northeastern Asia (Chernetsov, 1973). A.P. Okladnikov (1948) has demonstrated that the Neolithic people of Baikal maintained contacts with remote areas of Western Siberia and those west of the Urals. According to Okladnikov, ties with the Baikal Neolithic are present in Neolithic cultures of the Yenisei and are due to the spread of Early Neolithic Baikalians in the southern and western direction. In fact, the Kitoy culture displays especially close ties with remote groups

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Table 6. Distances between Eskimos, Chukchi, Aleuts, and ancient Chukotkans and southern groups Groups

Eskimos

Chukchi Coastal

Reindeer

Aleuts

Welen

Ekven

Ancient Chukotkan, pooled

Burmese

0.109

0.106

0.130

0.090

0.065

0.063

0.058

Indians

0.187

0.183

0.191

0.130

0.129

0.126

0.120

Malays

0.154

0.107

0.132

0.102

0.107

0.123

0.109

Australians

0.268

0.285

0.303

0.192

0.194

0.138

0.164

Papuans, Murua

0.105

0.088

0.113

0.077

0.057

0.073

0.059

Papuans, Awayama

0.122

0.132

0.145

0.098

0.084

0.082

0.078

Papuans, pooled

0.109

0.105

0.125

0.083

0.066

0.072

0.064

Melanesians

0.250

0.228

0.260

0.178

0.188

0.152

0.171

both in the east and in the west (Okladnikov, 1974). The Lake Baikal area appears to have been the source of migrations to northern Yakutia, where Neolithic sites are markedly similar to those of Baikal. The Yakutian Neolithic, in turn, inÀuenced the Neolithic of Chukotka, affecting the course of further cultural evolution in that region, including the emergence of the Chukchi, Koryak, and Itelmen cultures (Dikov, 1974). At the same time, the possibility of ancient migrations to Chukotka from the south, along the Pacific coast, cannot be ruled out. Certain writers pointed to the Paci¿c Asian route of the peopling of America via Beringia (Zubov, 2002; Neves et al., 2003). With this in view, we compared prehistoric Chukotkans and recent Eskimos, Chukchi, and Aleuts with southern Mongoloids, Indians, Australians, Papuans, and Melanesians (Table 6). Whereas none of the modern Paci¿c groups show any southern links, the ancient Chukotkans do display a certain similarity with modern Burmese and Papuans. Notably, the southern Pacific tendency of Welen and Ekven was also demonstrated using another battery of nonmetric cranial traits (Kozintsev, 1988). Therefore the admixture of migrants from inland Siberian populations with those from more southern areas of the Paci¿c Basin and the adaptation of both to the extreme environment of Chukotka were likely important factors in proto-Eskoaleut origins.

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Received September 1, 2011.

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