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Mtdna Diversity in Chukchi and Siberian Eskimos: Implications for the Genetic History of Ancient Beringia and the Peopling of the New World Yelena B

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Mtdna Diversity in Chukchi and Siberian Eskimos: Implications for the Genetic History of Ancient Beringia and the Peopling of the New World Yelena B Am. J. Hum. Genet. 63:1473±1491, 1998 mtDNA Diversity in Chukchi and Siberian Eskimos: Implications for the Genetic History of Ancient Beringia and the Peopling of the New World Yelena B. Starikovskaya,1 Rem I. Sukernik,1 Theodore G. Schurr,2,3 Andreas M. Kogelnik,2,3 and Douglas C. Wallace2,3 1Laboratory of Human Molecular Genetics, Institute of Cytology and Genetics, Siberian Division, Russian Academy of Sciences, Novosibirsk, and Chukotkan Research Center, Far Eastern Division, Russian Academy of Sciences, Anadyr, Chukotkan Autonomous Region; and 2Department of Anthropology and 3Center of Molecular Medicine, Emory University, Atlanta Summary Introduction The mtDNAs of 145 individuals representing the abo- Archaeological records of Kamchatka, Chukotka, and riginal populations of ChukotkaÐthe Chukchi and Si- Alaska that date back to 15,000 years before the present berian EskimosÐwere subjected to RFLP analysis and (YBP) suggest that the latest inhabitants of the Bering control-region sequencing. This analysis showed that the land bridge were living in rich unglaciated lowlands and core of the genetic makeup of the Chukchi and Siberian perhaps were occupying these areas of ancient Beringia Eskimos consisted of three (A, C, and D) of the four as two distinct populations. One was apparently an in- primary mtDNA haplotype groups (haplogroups) (A±D) land population of mammoth and bison hunters, observed in Native Americans, with haplogroup A being whereas the other consisted of small groups of ®shing the most prevalent in both Chukotkan populations. Two and sea mammal±hunting populations who were scat- unique haplotypes belonging to haplogroup G (formerly tered along the coast of southern Beringia. The sub- called ªotherº mtDNAs) were also observed in a few mergence of most of the Bering land bridge and subse- Chukchi, and these have apparently been acquired quent formation of the Bering Strait 10,000 YBP » through gene ¯ow from adjacent Kamchatka, where separated Chukotka and Alaska and may have affected haplogroup G is prevalent in the Koryak and Itel'men. the overall divergence of these groups, because of the In addition, a 16111CrT transition appears to delineate dislocation of interior and coastal populations (Young an ªAmericanº enclave of haplogroup A mtDNAs in 1988; Hoffecker et al. 1993; Dikov 1994; King and Slo- northeastern Siberia, whereas the 16192CrT transition bodin 1996). Eventually, a few distinct populations demarcates a ªnorthern Paci®c Rimº cluster within this evolved in the northern Paci®c Rim, and presumably haplogroup. Furthermore, the sequence-divergence es- these are re¯ected genetically in the Paleoasiatic-speak- timates for haplogroups A, C, and D of Siberian and ing Chukchi, Na-Dene±speaking Indians, and Eskimo- Native American populations indicate that the earliest Aleuts (Krauss 1988). The aboriginal inhabitants of inhabitants of Beringia possessed a limited number of ChukotkaÐthe Chukchi and Siberian EskimosÐare founding mtDNA haplotypes and that the ®rst humans likely the survivors of these rapid environmental changes expanded into the New World 34,000 years before and, hence, are of great importance for understanding » present (YBP). Subsequent migration 16,000±13,000 the evolution of both Siberian and Native American YBP apparently brought a restricted number of haplo- populations. group B haplotypes to the Americas. For millennia, Ber- The utility of the unique genetic properties of mtDNA, ingia may have been the repository of the respective including its maternal inheritance and high mutation founding sequences that selectively penetrated into rate, have been helpful in addressing long-standing ques- northern North America from western Alaska. tions about the timing and origins of the ®rst Americans, the number of subsequent demic expansions across Ber- ingia, the chronology and routes of these expansions, and the sizes of the founding populations (Wallace et al. Received May 15, 1998; accepted for publication August 19, 1998; 1985; Ward et al. 1991; Neel et al. 1994; Wallace 1995; electronically published October 27, 1998. Forster et al. 1996). Our ®rst survey of mtDNA variation Address for correspondence and reprints: Dr. Douglas C. Wallace, in the Chukchi and Siberian Eskimos involved partial Director, Center for Molecular Medicine, Emory University School of haplotype analysis and showed that the ªreindeerº Medicine, 1462 Clifton Road N.E., Atlanta, GA 30322. E-mail: Chukchi exhibited three (A, C, and D) of the four [email protected] q 1998 by The American Society of Human Genetics. All rights reserved. mtDNA haplogroups (A±D) observed in Native Amer- Lys 0002-9297/98/6305-0027$02.00 icans but lacked the COII/tRNA intergenic 9-bp de- 1473 1474 Am. J. Hum. Genet. 63:1473±1491, 1998 letion associated with haplogroup B (Torroni et al. mtDNAs merit additional analysis and that such data 1993a, 1993b). In contrast, Siberian Eskimos showed would also be extremely helpful in the reconstruction of mtDNAs from only two of these haplogroups (A and the evolutionary history of Beringia. In the present study, D), and appeared to be the only aboriginal Siberian we have further characterized the mtDNA variation in group lacking ªotherº haplotypes2that is, mtDNAs that Chukotkan populations, via high-resolution restriction are not within haplogroups A±D (Torroni et al. 1993b; analysis and CR sequencing, and have attempted to trace Sukernik et al. 1996). When native Siberian mtDNAs their origins by comparing the resulting RFLP haplo- were subjected to high-resolution restriction analysis, types and CR lineages with those present in other Si- most of these ªotherº haplotypes were shown to be eth- berian and Native American populations. nic-, tribal-, or region-speci®c haplotypes that clustered into additional haplogroups. This was seen most clearly Subjects and Methods in the Udegeys of the Sikhote Alin and in the Nivkhs of the lower Amur/Sakhalin Island region (Torroni et al. Populations 1993b). Similarly, previous analyses of restriction-site varia- Chukchi.ÐWhen the ®rst Russians reached the mouth tion in Native American mtDNAs had revealed that of the Anadyr River in 1648, the Paleoasiatic-speaking nearly all of these mtDNAs clustered within haplogroups Chukchi were reindeer and mountain-sheep hunters who A±D. Each of these haplogroups was widespread, al- wandered within speci®c hunting areas in eastern Chu- though they were unevenly distributed among the dif- kotka and numbered !2,000 individuals (Dolgikh ferent tribes and linguistic groups in the New World, 1960). By the end of the 19th century, shortly after the and each haplogroup traced back to only one or two transition from a hunting-®shing subsistence to a rein- founding mtDNA haplotypes that had originated in Asia deer economy, the Chukchi had considerably increased (Wallace 1985; Schurr et al. 1990; Torroni et al. 1992, in their population size and had spread widely in Chu- 1993a). As a result, the variation that had accumulated kotka, pushing and assimilating adjoining Yukagir tribes within each haplogroup since the ®rst humans arrived in the west, the Koryaks living south of the lower Anadyr in the Americas was quanti®ed and was found to give region, and the Eskimos residing along the entire coast divergence times, for haplogroups A, C, and D, of of the Chukchi peninsula (Gondatti 1897; Bogoras 34,000±26,000 YBP. Since haplogroup B was not found 1910). With the passage of time, some Chukchi families in eastern Siberia but was prevalent in the Americas, it (present-day coast Chukchi) settled on the coast and has been proposed that haplogroup B represents an in- adopted a way of life similar to that of Siberian Eskimos dependent population expansion into the New World living in adjoining villages (Bogoras 1910). (Torroni et al. 1992, 1993b, 1994b). Siberian Eskimos.ÐBy the turn of the 20th century, However, the results of other studies are at odds with the territory inhabited by Siberian Eskimos had been this interpretation of a limited number of Asian founding reduced to three small enclaves on the northeastern and mtDNA lineages populating the Americas and an southern coast of the Chukchi peninsula. These were ªearlyº entry into the New World (Ward et al. 1991, populated by three Yupik-speaking tribesÐthe Naukan, 1993; Shields et al. 1993; Merriwether et al. 1995). On Chaplin, and Sireniki Eskimos (Rudenko 1947; Men- the basis of the low diversity and small mean pairwise ovshchikov 1959; Arutiunov et al. 1982; Krauss 1994). sequence differences of mtDNA noncoding control re- At that time, Naukan Eskimos inhabited the mountain gion (CR) sequences within and between Chukchi, Si- terrace located southeast of Uelen, the settlement of berian Eskimos, Alaskan Na-Dene Indians, and Alaskan coastal Chukchi. They spoke their own dialect of Si- and western Greenland Eskimos, Shields et al. (1993) berian Yupik and occasionally intermarried with the estimated that the average date for the ancestry of these nearby Eskimo tribe of Little Diomide Island (®g. 1). mtDNA lineages is 7,100±5,100 YBP. In contrast, the The Chaplin Eskimos were living in a number of settle- mean pairwise sequence differences within the Amer- ments scattered along the coastline between Arakam- indian tribesÐthe Nuu-Chah-Nulth and the Ya- chechen Island and Provideniya (Plover) Bay. They were kimaÐgave estimated date of 13,200±12,100 YBP, im- isolated from the Naukan Eskimos by geographic dis- plying a ªlateº entry of ancient Native Americans into tance, and Chukchi settlements were interspersed be- the New World (Shields et al. 1993). However, the small tween them. Marital exchanges between Chaplin Eski- sample sizes of the Chukchi and of the Siberian and mos and St. Lawrence Island Eskimos, who spoke the Alaskan Eskimos (seven, six, and ®ve individuals, re- same dialect of Siberian Yupik, were common in his- spectively) that Shields et al. (1993) used for the analysis torical times (Menovshchikov 1959). The Sireniki tribal of CR sequences could have in¯uenced these results group lived in a few settlements located west of Provi- considerably.
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