Mitochondrial DNA of Protohistoric Remains of an Arikara Population from South Dakota: Implications for the Macro- Siouan Language Hypothesis
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Mitochondrial DNA of Protohistoric Remains of an Arikara Population from South Dakota: Implications for the Macro- Siouan Language Hypothesis Diana M. Lawrence,1 Brian M. Kemp,2 Jason Eshleman,3 Richard L. Jantz,4 Meradeth Snow,3 Debra George,3 and David Glenn Smith3 Abstract Mitochondrial DNA (mtDNA) was extracted from skeletal re- mains excavated from three Arikara sites in South Dakota occupied between AD 1600 and 1832. The diagnostic markers of four mtDNA haplogroups to which most Native Americans belong (A, B, C, and D) were successfully identified in the extracts of 55 (87%) of the 63 samples studied. The frequen- cies of the four haplogroups were 42%, 29%, 22%, and 7%, respectively, and principal coordinates analysis and Fisher’s exact tests were conducted to compare these haplogroup frequencies with those from other populations. Both analyses showed closer similarity among the Mohawk, Arikara, and Sioux populations than between any of these three and any other of the com- parison populations. Portions of the first hypervariable segment (HVSI) of the mitochondrial genome were successfully amplified and sequenced for 42 of these 55 samples, and haplotype networks were constructed for two of the four haplogroups. The sharing of highly derived lineages suggests that some recent admixture of the Arikara with Algonquian-speaking and Siouan- speaking groups has occurred. The Arikara shared more ancient lineages with both Siouan and Cherokee populations than with any other population, con- sistent with the Macro-Siouan language hypothesis that Iroquoian, Siouan, and Caddoan languages share a relatively recent common ancestry. The popular concept of Plains Indians is dominated by the equestrian nomads, such as the Dakota, most recently popularized in the film Dances with Wolves. However, before the 19th century, semisedentary tribes living in earth lodge vil- lages were the primary inhabitants of the Great Plains. The Arikara, or Sahnish, 1Forensic Analytical Sciences, Hayward, CA 94545. 2Department of Anthropology and the School of Biological Sciences, Washington State University, Pullman, WA 99164. 3Molecular Anthropology Laboratory, Department of Anthropology, University of California, Davis, CA 95616. 4Department of Anthropology, University of Tennessee, Knoxville, TN 37996. Human Biology, April 2010, v. 82, no. 2, pp. 157–178. Copyright © 2010 Wayne State University Press, Detroit, Michigan 48201-1309 key words: mtDNA haplogroup, HVSI haplotype, ancient DNA, migra- tion, admixture, Arikara, Mohawk, Sioux, Chippewa, Cherokee, Ojibwa, Pawnee, Algonquian groups, South Dakota. 158 / lawrence et al. living along the banks of the Missouri River in South Dakota, were one of the most populous and powerful tribes to emerge into the historic period. They grew corn, beans, and squash in the fertile soils along the Missouri floodplain while continuing to follow a seasonal hunter-gatherer lifestyle collecting wild plants and hunting bison (Blakeslee 1994). This subsistence pattern was unique in North America in its heavy reliance on hunting. Lehmer and Wood (1977) have likened it to the mixed horticultural and pastoral economies of the Old World. Arikara is a Caddoan language, closely related to Pawnee (Parks 1979), from which it split between 1450 and 1650 (Parks 1979). Linguistic relatives far- ther south include Wichita, Kitsai, and Caddo proper (Parks 1979). The sites in South Dakota attributed to the Arikara represent the northernmost extension of the Caddoan languages until the group moved into North Dakota after leaving the Leavenworth site in 1832. Because the focus of our study is the relationship of the language spoken by the Sahnish to other languages, we have chosen to use Arikara to refer to the Sahnish throughout this paper. Archaeological complexes usually regarded as directly ancestral to the his- toric Arikara include the Extended Coalescent Variant (ca. AD 1400/1450–1650), the Postcontact Coalescent (ca. 1650–1785) (Johnson 1998), and the Disorganized Coalescent (1785–1862) (Lehmer 1971). Arikara connections to early archaeologi- cal complexes, such as the Initial Coalescent (1300–1600) or the earlier Central Plains Tradition (900–1450) in Kansas and Nebraska, have not been established, al- though some sort of relationship has been widely assumed. If there is a connection, it likely predates the emergence of the Arikara as an ethnic entity. Exposure of the Arikara to Europeans resulted in dramatic changes, which affected their health and well-being as well as their population structure. Initial European contact may have happened as early as 1700 but is not documented before about 1706 (Jantz and Owsley 1994). Initially, European contact was bene- ficial to the Arikara. Acquisition of the horse increased bison hunting efficiency, and the Arikara eventually established themselves as middlemen in trade rela- tions, which initially brought a wave of prosperity but ultimately resulted in al- most complete cultural collapse as epidemics of European diseases decimated their numbers. During the early postcontact period, the Arikara population was estimated to be about 30,000 individuals (Holder 1970: 30). By the early 19th century this number had been reduced to the approximately 2,000 individuals living in the vil- lages along the Grand River in South Dakota, now known as the Leavenworth site. Shortly after leaving the Grand River villages, the Arikara moved their village to Ft. Clark, North Dakota, where their numbers were further reduced by a par- ticularly virulent epidemic sweeping Plains tribes in 1837–1838 (Trimble 1994). In 1862 the Arikara, Mandan, and Hidatsa were moved to the Fort Berthold In- dian Reservation, where they now live as the Three Affiliated Tribes. The Arikara population nadir was 380 individuals, reported by the 1904 census, but by 2005 they had increased to 3,400, as reported by the Indian Health Service. Today the Three Affiliated Tribes lead lives much like those of many other Native American mtDNA of Protohistoric Arikara / 159 tribes. They operate a casino, encourage tourism, and are attempting to develop the natural resources on their lands, particularly oil and gas. Chafe (1976) proposed that Caddoan together with the Iroquoian and Siouan languages belongs to a larger language family that he called Macro-Siouan, whose origin was in the Southeast. Previous studies have indicated that populations liv- ing in the same geographic area or closely related to each other often exhibit similar mitochondrial DNA (mtDNA) haplogroup frequencies as a result of either common ancestry and/or admixture (Bolnick and Smith 2003). Although the speakers of related languages themselves need not be closely genetically related to each other, they often are when vertical transmission of language predominates, and thus relationships between languages provide testable genetic hypotheses. To date, comparative studies of the mtDNA of the Caddoan, Iroquoian, and Siouan groups have not been conducted. The Iroquoian languages are divided into two major branches: Northern Iroquoian, which includes Mohawk, and Southern Iroquoian, which includes Cherokee (Chafe 1976). The Northern Iroquoian peoples split from their south- ern counterparts and moved north sometime between 3,500 and 4,000 years ago (Snow 1994). Subsequent admixture with Algonquian tribes in neighboring areas might make their genetic structure resemble a hypothetical admixture between ex- tant Cherokee and Algonquian tribes. If historic Plains tribes, such as the Arikara and the Sioux, also admixed with neighboring Algonquian tribes after migrating from their hypothetical southeastern homeland, then the genetic structures of the Arikara, Mohawk, and Sioux might resemble each other but differ from those of Cherokee groups. The ancestors of all three tribes might have more closely resembled the Cherokee before migrating northward, where they encountered and may have mixed with Algonquian groups. The identification of defining mtDNA markers, using restriction analysis or sequencing, makes it possible to assign individuals in populations to their hap- logroups or subhaplogroups. Additional markers or DNA sequences can further subdivide members of each haplogroup, or subhaplogroup, into different haplo- types, each defined by a unique set of genetic markers or nucleotide sequences. Comparisons of numbers of nucleotide differences between all paired sequences in a lineage or population provide an estimate of the extent of diversity among in- dividuals within that lineage or population. Because most nonbasal (i.e., derived) mtDNA haplotypes are tribe specific or nearly so (Malhi et al. 2002), the sharing of the same or similar haplotypes reflects either shared ancestry or admixture among different populations that are geographically proximate (Schurr 2004). The frequency distributions of haplogroups can be used to genetically characterize populations. These distributions usually exhibit geographic struc- ture, because the probability that mating occurs between members of any two different human populations is inversely correlated with the geographic distance separating them. Thus the pattern of genetic diversity within and between human populations is sometimes consistent with a model of isolation by distance. The mtDNA of modern Native Americans has been shown to fall into one of at least 160 / lawrence et al. five haplogroups: A, B, C, D, and X (Kaestle and Smith 2001; Malhi et al. 2001, 2002). These haplogroups are defined by diagnostic mutations (O’Rourke et