24
49 CHAPTER 4 A Land of Diversity: Genetic Insights into Ancestral Origins John R. Johnson, Brian M. Kemp, Cara Monroe, and Joseph G. Lorenz Native California has long been recognized for its high degree of linguistic diversity. At least 78 mutually unintelligible languages have been documented for native peoples who lived within the boundaries of the current state of California and along the Baja California peninsula (Goddard 1996; Golla 2011; Laylander 1997; Mithun 1999). The mul- tiplicity of languages present in this region is generally derived from past migrations and cultural adaptation to a diverse range of ecological types (Golla 2007; Moratto 1984; Nichols 1992). Reconstructing the record of ancient migrations has been approached through techniques of his- torical linguistics, archaeology, oral tradition, morphometric measure- ments of skeletal anatomy, and most recently by DNA studies (Breschini 1983; Bright and Bright 1976; Eshleman et al. 2004; Johnson and Lorenz 2006; Laylander 2010; Levy 1997; Sutton 2009). Genetic stud- ies, in particular, offer a direct means of testing hypotheses constructed by archaeologists and linguists regarding prehistoric population move- ments that produced specific mosaics of languages and cultures present in aboriginal California at the time of European contact. The discipline of molecular anthropology is relatively new, yet it already is beginning to contribute important insights to the study of California prehistory. UNDERSTANDING THE BASICS OF DNA ANALYSIS Before discussing the application of genetic research to understanding ancient migratory patterns, some background information is necessary. As everyone is taught in introductory biology courses, humans inherit half of their 46 chromosomes maternally and the other half paternally. Contemporary Issues in California Archaeology, edited by Terry L Jones and Jennifer E. Perry, 49–72. ©2012 Left Coast Press Inc. All rights reserved.

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Chapter 4

a Land of Diversity: Genetic Insights into ancestral Origins

John R. Johnson, Brian M. Kemp, Cara Monroe, and Joseph G. Lorenz

Native California has long been recognized for its high degree of linguistic diversity. At least 78 mutually unintelligible languages have been documented for native peoples who lived within the boundaries of the current state of California and along the Baja California peninsula (Goddard 1996; Golla 2011; Laylander 1997; Mithun 1999). The mul-tiplicity of languages present in this region is generally derived from past migrations and cultural adaptation to a diverse range of ecological types (Golla 2007; Moratto 1984; Nichols 1992). Reconstructing the record of ancient migrations has been approached through techniques of his-torical linguistics, archaeology, oral tradition, morphometric measure-ments of skeletal anatomy, and most recently by DNA studies (Breschini 1983; Bright and Bright 1976; Eshleman et al. 2004; Johnson and Lorenz 2006; Laylander 2010; Levy 1997; Sutton 2009). Genetic stud-ies, in particular, offer a direct means of testing hypotheses constructed by archaeologists and linguists regarding prehistoric population move-ments that produced specific mosaics of languages and cultures present in aboriginal California at the time of European contact. The discipline of molecular anthropology is relatively new, yet it already is beginning to contribute important insights to the study of California prehistory.

Understanding the Basics of dna analysis

Before discussing the application of genetic research to understanding ancient migratory patterns, some background information is necessary. As everyone is taught in introductory biology courses, humans inherit half of their 46 chromosomes maternally and the other half paternally.

Contemporary Issues in California Archaeology, edited by Terry L Jones and Jennifer E. Perry, 49–72. ©2012 Left Coast Press Inc. All rights reserved.

50 |

John R. Johnson, Brian M. Kemp, Cara Monroe, and Joseph G. Lorenz

The nucleus of each cell holds these chromosomes that are composed of four nucleotide bases (adenine, guanine, cytosine, and thymine), the order of which codes for a person’s biologically inherited characteristics. In total, the human genome comprises approximately three billion nucle-otides. Outside the nucleus are found hundreds of mitochondria, the cell’s “energy factories,” that each contain a number of their own small circular genomes. The first human mitochondrial genome was sequenced in 1981, which totaled only 16,569 nucleotides in length (Anderson et al. 1981; Andrews et al. 1999). Unlike the nuclear DNA, mitochondrial DNA is inherited only maternally and thus presents an unmixed record of a per-son’s direct female ancestry. Both nuclear and mitochondrial DNAs are composed not only of genes that are subject to natural selection, but also of noncoding segments that can accumulate mutations (genetic markers) over many generations and thus provide a record of an individual’s ances-try. The variability of mutations among a large number of individuals can be compared and contrasted to understand ancestral relationships between different peoples, thus providing clues to migratory patterns of human populations as they spread throughout the world.

To date, nearly all California Indian genetic studies have been based upon mitochondrial DNA (mtDNA) analysis. Studies of genetic markers in the nuclear genome, particularly those found on the Y chromosome, which is inherited only by males, can produce important information (e.g., Schroeder et al. 2009; Zegura et al. 2009); however, mtDNA is more abundant, relatively easy to analyze, and more likely to be pre-served in ancient remains. Because mtDNA is maternally inherited, it is unaffected by European male gene flow and has survived in greater measure among living California Indian descendants due to differential marriage patterns and mortality rates of men and women after missioni-zation. Thus, mtDNA analyses have had a considerable head start in producing our first glimpses into genetic patterns that existed among the diverse peoples of aboriginal California prior to European contact. This is thus far ideal, as mtDNA diversity has been extensively documented among indigenous American populations. In addition, the mutation rate or “molecular clock” in mtDNA has been useful in identifying and dat-ing initial colonization events and in documenting migration.

Worldwide, human mtDNAs have been divided into haplogroups, groups of maternal lineages that exhibit mutations in common, denot-ing a common origin at some time in the past. The nomenclature for each identified haplogroup has been assigned according to letters of the alphabet. Five indigenous mtDNA haplogroups characterize living Native Americans (Merriwether 2006; Torroni 2000), designated A, B, C, D, and X (X being the least common and only reported twice among 425 California Indian mtDNAs sampled to date, ancient and modern). Following the capital letter assigned to each haplogroup,

A Land of Diversity: Genetic Insights into Ancestral Origins |

51

numbers and lowercase letters are used for subgroups. For example, D4h3a is the official designation provided for a particular clade (daugh-ter lineage) within mitochondrial haplogroup D that has been identified in very ancient remains from Alaska, but thus far have only been found among Chumash peoples of all living Native Americans north of Mexico (Johnson and Lorenz 2006; Kemp et al. 2007; Perego et al. 2008).

A comparison of mtDNA haplogroup distributions among popula-tions in different regions can reveal similarities that reflect ancestral relationships; however, such similarities can also be misleading. A more refined means of demonstrating ancient connections between popula-tions, as well as determining the age of a particular clade, occurs at the haplotype level of analysis. Haplotypes are specific lineages within a haplogroup, identified by additional mutational variation beyond that defining the haplogroup. To date, all analyses of California Indian haplo-types (or specific lineages) have been based on comparing mutations that occur in two hypervariable, noncoding segments of the mtDNA mol-ecule. Commonly, only a portion of Hypervariable Segment I (HVSI), numbered between nucleotide positions (np) 16051 and 16365 of the mtDNA molecule, has been used in comparative analyses of California Indian mitochondrial genomes (e.g., Johnson and Lorenz 2006; Lorenz and Smith 1997). The remain portions of the d-loop, including Hypervariable Segment II (HVSII), have not yet been analyzed to any significant extent among California Indian populations, although it has yielded important discoveries for understanding prehistoric population expansions in adjacent regions (e.g., Kemp et al. 2010).

Advancing technology has facilitated full mitochondrial genome scans that identify additional genetic markers outside of the two mtDNA hypervariable segments (HVSI and HVSII). Such studies have advanced our understanding of the ancestral links of Native Americans to popu-lations in Asia and the number of founding haplotypes (lineages) that arrived from Beringia during the initial peopling of the Americas (Achilli et al. 2008; Fagundes et al. 2008; Kemp and Schurr 2010; Tamm et al. 2007). Recently, complete mitochondrial genomes have been collected for California Indian descendants, but the number of samples analyzed thus far is still too small for useful comparative analysis. Also, given the problems of degradation and contamination of DNA molecules in ancient bone, it is practical and cost-effective to obtain HVSI sequences than to attain complete genome results. Thus HVSI sequence variabil-ity, rather than full mtDNA genome data, forms the basis of the fol-lowing analyses of population differentiation as well as dating the divergence of lineage or haplotype clades. For depicting phylogenetic relationships among haplotypes, network diagrams become useful (see Figures 4.3–4.6 for examples), wherein the daughter lineages radiate from a central, founding haplotype (Bandelt et al. 1999).

52 |

John R. Johnson, Brian M. Kemp, Cara Monroe, and Joseph G. Lorenz

genetic and lingUistic stUdy of california’s Prehistoric Migrations

Although genetic and cultural variables can diverge independently, there are many instances, however, where genes, language, and cul-ture coevolve. Certainly, populations who migrate from one location to another also bring their native language and customs. These cultural links can be maintained if connections with their ancestral territory are maintained. If not, the language and culture of the emigrating group will begin to differentiate, based in part upon the new natural and social envi-ronmental conditions encountered. If the territory into which an incom-ing group arrives was already inhabited, either population replacement (through hostile action) or intermarriage can occur. Over time, culture and language may be so altered that any connections with the region of origin is obscured. Despite this, genetic data provide particularly useful clues for reconstructing population histories and provide an independent yet complementary means of detecting past migrations.

Directly testing various migration scenarios through ancient DNA (aDNA) research presents an even more powerful method of determin-ing population prehistory, as ancient demographic events such as migra-tion and intermarriage (gene flow), genetic drift (isolation), or admixture may be obscured by recent population history. Distinguishing migratory events can be difficult to identify with modern DNA data alone. Genetic data from prehistoric populations in combination with archaeological data (such as radiocarbon dating) can, therefore, reveal aspects of unrec-ognized population history such as lineage extinction and replacement.

An overview of California Indian linguistic prehistory by Golla (2007, 2011) analyzed more than a century of linguistic research and identified approximately seven migration events that resulted in language family spreads. The Hokan stock (superfamily) is composed of five language families (Shastan, Palaihnihan, Pomoan, Yanan, and Yuman-Cochimi), plus five isolate languages (Karuk, Chimariko, Washo, Esselen, and Salinan). The Penutian stock (superfamily) in California is represented by five language families (Plateau Penutian, Wintuan, Maiduan, Utian, and Yokutsan). Not related to the Hokan and Penutian macro-units are five additional, unrelated language families: Algic, Athapaskan, Yukian, Chumashan, and Uto-Aztecan (Goddard 1996; Golla 2007; Mithun 1999).

Yukian and Chumashan have been hypothesized to be the oldest lan-guage families in Alta California, possibly descended from the earliest Paleoindian entrants into the region (Golla 2011). The diverse languages and language families that have been included within the Hokan stock are widely scattered around California and are considered to have been

A Land of Diversity: Genetic Insights into Ancestral Origins |

53

part of population movements that occurred during the Early Holocene. The language families associated with the Penutian stock represent multiple migrations that began later during Middle Holocene times. Yokutsan and Utian, two closely related languages within the Penutian linguistic group, are thought to have been brought to California by some of the earliest emigrants from either the Great Basin or Plateau regions, whereas peoples speaking Wintuan and Maiduan languages arrived during the Late Holocene. The Wiyot and Yurok languages may have arisen originally in the Columbian Plateau region and then spread to the coast, before speakers of these languages were pushed southward into northern California by the Late Holocene expansion of Athapaskan-speaking peoples (Golla 2007). Most linguists favor a homeland for Uto-Aztecan (Takic) languages in the greater Southwest with an expan-sion into California, perhaps first into the southern Sierra Nevada and adjacent San Joaquin Valley, during the early part of the Late Holocene. Subsequent migrations resulted in the extension of the Numic and Takic branches into the Great Basin and southern California, respectively, dur-ing the Late Holocene (Laylander 2010; Moratto 1984; Sutton 2009).

These various linguistic scenarios can be tested using mtDNA evidence. Comparative analysis of haplogroup and haplotype frequen-cies and distributions between the hypothesized regions of origin and particular California ethnolinguistic groupings were used to test ances-tral relationships. Dating of mtDNA lineages will use two sources: (1) mtDNA samples contributed by modern California Indian descend-ants with documented genealogies (Table 4.1) and (2) ancient DNA derived from prehistoric skeletal remains (Table 4.2). Within a mito-chondrial haplogroup, the age of the ancestral node for a particular clade can be estimated by using a calculated rate of mutation (Kemp et al. 2007), whereas in the case of ancient samples, direct radiocarbon dating can be paired with DNA findings.

california indian Mtdna Patterns

A total of 207 samples contributed by living California Indian descend-ants and 218 samples reported from aDNA studies are presented in Tables 4.1 and 4.2. Table 4.1 includes all but two of the 126 matrilines (maternal lineages) reported by Johnson and Lorenz (2006), as well as 17 newly contributed samples (on file at the Santa Barbara Museum of Natural History, Santa Barbara), 33 Washo and Northern Hokan sam-ples reported by Kaestle (1998), 8 matrilines identified by Schroeder et al. (2011), and 25 Baja California samples studied by Monroe et al. (2008). It should be noted that most of the modern samples reported here refer to the number of sampled matrilines that have been determined to have

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A Land of Diversity: Genetic Insights into Ancestral Origins |

59

been independent of each other based upon genealogical research using archival records. For the remainder (particularly the Washo, Pomo, Karuk, and most Baja California samples), the same level of genea-logical investigation was not undertaken; however, at least for the Baja California samples, it was determined that the matrilines were independ-ent back to the maternal grandmother.

Table 4.2 presents data from both published and unpublished reports pertaining to aDNA findings and associated radiocarbon dates. More than half of the aDNA samples come from studies conducted in the San Francisco Bay region, the largest portion (71 samples) from CA-SCL-38, the Yukisma Site (Monroe et al. 2011; Villanea 2010).

A comparison of haplogroup distributions among living populations is instructive, sometimes reflecting linguistic differences and sometimes regional differences (Figure 4.1). Among Yuman (Southern Hokan) peo-ples speaking Diegueño (Ipai, Kumeyaay, and Tipai) and Cochimí lan-guages, only haplogroups B and C have been reported to date. Peoples speaking Uto-Aztecan languages in the California region also exhibit high frequencies of haplogroup B and C matrilines, with haplogroup D an important minority component in some groups. Significantly, this pattern of high frequencies of haplogroup B and C is predominant in the American Southwest (Kemp et al. 2010).

Haplogroup A is almost entirely absent among Uto-Aztecan groups in California, except for two Luiseño matrilines. Chumash peoples in the Santa Barbara Channel area reflect quite different genetic lineages compared with that of their Uto-Aztecan neighbors, with haplogroups A and D predominating. The distribution of haplogroup A also extends up the coast into the Esselen and Salinan populations; however, the presence of haplogroups B and D may be consistent with intermarriage between these Hokan groups and their Costanoan and Yokuts neighbors. Penutian groups of the San Francisco Bay area and Central Valley in general exhibit high frequencies of haplogroups B and D. For groups inhabiting the Coast Ranges north of San Francisco, only limited data are available, but those samples reported in Table 4.1, along with an unpublished study by Schurr et al. (2011), appear to indicate that matrilines belonging to haplogroup B are quite common in this region, whereas haplogroup C is rare and haplogroup A is completely absent. The only Hokan-affiliated enclave in the Great Basin region, the Washo group indigenous to the Lake Tahoe area, consists mostly of haplogroups B and C (Kaestle 1998).

The haplogroup distributions derived from living California Indians and their descendants can be compared with aDNA findings to gain insights regarding genetic continuity and the appearance of new line-ages that might reflect immigrant contributions to the local gene pool, or conversely lineage extinction.

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In southern California, two patterns emerge from the limited aDNA data generated to date. When Uto-Aztecan peoples initially arrived in California is hotly debated and of considerable interest to regional scholars (for recent summaries of research on this topic, see Laylander 2010; Sutton 2009). Interestingly, the limited data avail-able from three inland sites dating to the Middle and Middle/Late Transition periods largely exhibit haplogroup distributions similar to descendants of Takic groups who inhabited the area at the time of European contact. For the Southern Channel Islands, homeland of the Island Gabrielino, a somewhat different picture emerges, with haplo-group A more common in prehistoric samples than among mainland Takic peoples. As has been suggested by Potter and White (2009), this may reflect intermarriage with Chumash inhabitants of the Northern Channel Islands and/or a genetic mixing as a new Gabrielino-speaking population moved to the southern islands. Increased sampling and/or analysis of these samples at the haplotypic level could add resolution to this supposition.

Figure 4.1 Mitochondrial haplogroup percentages for ethnolinguistic groups and ancient sites.

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For the Chumash region, the ancient samples accumulated to date nearly all derive from the Northern Channel Islands. With the exception of one haplogroup C lineage (which also occurs in a minority of surviv-ing Chumash matrilines), all ancient samples belong to haplogroup A. The specific haplotypes present among ancient Santa Cruz Island sam-ples, when compared with those reported from living descendants by Johnson and Lorenz (2006), clearly demonstrate that Chumash ances-tors were inhabiting the Northern Channel Islands by 5,000 years ago (Monroe et al. 2010). Although few ancient samples are available for the Central Coast north of the area inhabited by Chumashan speakers, a preliminary analysis by Breschini and Haversat (2008a) suggests that haplogroup A might well have been common among Esselen speakers. They hypothesized that two haplogroup D samples that were obtained from burials during the latest occupation of CA-MNT-831 in Monterey represent incoming Costanoans, who surrounded Monterey Bay at the time of Spanish contact. A more fine-scale analysis at the haplotypic level would permit a better test of this notion.

The ancient DNA results from the San Francisco Bay area and the Central Valley resemble the haplogroup distribution found among mod-ern Penutian peoples of the region. However, the ancient distribution differs from modern samples; haplogroup C is in relatively high fre-quency in prehistoric burial populations, whereas the presence of haplo-group C among living Penutian groups is limited to four samples found among San Francisco Bay area peoples (three Costanoans and one Bay Miwok) and a single sample from the southernmost Yokuts tribe, the Hometwoli. More research will be necessary to determine if the rarity of haplogroup C among living descendants reflects a problem of sampling, or whether this haplogroup might have been more widely present among Hokan groups that became displaced or absorbed by incoming Penutian migrations. The high percentage of haplogroup C lineages among the Washo and Yuman-Cochimi peoples, all speaking languages classified as Hokan, suggests that haplogroup C lineages could well have been associated with the spread of Hokan languages during California’s Early Period; however, analysis at the haplotype level is needed to resolve this.

external relationshiPs

To investigate whether particular California groups are similar in their haplogroup distributions to other regions, we assembled a database of reported ancient and contemporary samples from adjacent regions. Aggregated haplogroup distributions for the Pacific Northwest, Plateau, and Great Basin, derived from the research of a number of published stud-ies, are shown in Figure 4.1 (Kaestle and Smith 2001; Malhi et al. 2004;

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Shields et al. 1993; Torroni et al. 1993; Ward et al. 1993). Comparisons were also made with populations from the American Southwest (Kemp et al. 2010; Malhi et al. 2003). A PCoA (principal coordinate analysis) plot of genetic distances,1 based on haplogroup frequencies, was con-structed to depict the relationship of California Indian groups to popu-lations in the other regions. The Yokuts haplogroup distribution was closest to ancient and modern Great Basin groups, and Takic (Southern California Uto-Aztecan) and Washo grouped with the ancient and mod-ern Southwest samples, whereas Chumash samples were isolated midway between Great Basin and Northwest Coast groups (Figure 4.2). These patterns to a certain extent support expectations based on previously presented hypotheses. The hypothesized Proto-Yokuts homeland of the ancient Great Basin fits with the high percentage of haplogroups B and D that they share with that region. Similarly, the spread of Uto-Aztecan languages into southern California from its presumed hearth among

Figure 4.2 Principal coordinates analysis based on haplogroup frequencies.

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agricultural populations of the American Southwest matches the findings from comparable haplogroup distributions between the two regions. The Washo’s similarity with Takic and Southwestern groups is less likely to be derived from common ancestral origins. Rather, this group’s lineages may represent a surviving enclave of an ancestral Hokan haplogroup dis-tribution that once was typical of Central California prior to the entry of groups speaking languages belonging to the Penutian stock. The genetic distinctiveness of Chumash peoples was anticipated, for Chumashan represents a linguistic isolate and there is a continuous archaeological record that reflects in situ cultural development over many millennia.

Phylogenetic differentiation and PoPUlation sPreads

The comparative study of haplogroup distributions, although insightful, is not as definitive as comparing specific haplotypes and determining ancestral relationships between these lineages within each haplogroup. Johnson and Lorenz (2006) produced network diagrams for the four principal haplogroups using HVSI sequence data for 121 samples (Figures 4.3–4.6). The network diagrams for mtDNA haplogroups

Figure 4.3 Haplogroup A network diagram for California Indian mtDNA lineages based on HVSI sequences (Johnson and Lorenz 2006). The 16093 “Chumash” clade is circled.

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Figure 4.4 Haplogroup B network diagram for California Indian mtDNA lineages based on HVSI sequences (Johnson and Lorenz 2006). The 16184 “Yok-Utian” and 16111 “Southwest-B2a” clades are circled.

Figure 4.5 Haplogroup C network diagram for California Indian mtDNA lineages based on HVSI sequences (Johnson and Lorenz 2006).

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A, B, and C each exhibit star-like patterns with a central, basal hap-lotype (founding lineage) and radiating daughter lineages or clades. Such star-like patterns typify expansion events; in this case, the basal haplotype represents the founding type brought into the Americas by female Paleoindian ancestors who migrated southward from Beringia. Most of the samples typed to haplogroup D also exhibit a star-like pat-tern, but one particular haplotype, four mutations removed in its HVSI sequence, actually represents a second founding haplotype within this haplogroup (arbitrarily designated D04 in Figure 4.6). This rare hap-lotype, now formally designated D4h3a, characterizes five Chumash matrilines. Its broader distribution, primarily found in both ancient and existing native peoples widely scattered along the western margin of the American continents, is consistent with a coastal migrating group that moved southward from Beringia (Johnson and Lorenz 2006; Kemp et al. 2007; Perego et al. 2008; Tamm et al. 2007).

Haplogroup A, in addition to exhibiting the star-like pattern char-acteristic for other mitochondrial haplogroups found among Native Americans, also exhibits a clade found only among Chumash groups in the form of a branching chain of haplotypes differentiated from the basal

Figure 4.6 Haplogroup D network diagram for California Indian mtDNA lineages based on HVSI sequences (Johnson and Lorenz 2006). The D4h3a haplotype has been shown to represent a separate founding lineage within haplogroup D (Kemp et al. 2007).

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haplotype by the presence of a C→T transition at np16093 (Figure 4.3). This specific mutation within the HVSI region of mtDNA haplogroup A has not been reported elsewhere among indigenous peoples in Western North America. Because other mutations followed the initial C→T tran-sition at np16093 and became fixed in descendant Chumash popula-tions, many millennia are implied for this group in the Santa Barbara region. Using a 34 percent mtDNA evolution for HVSI (taken as one estimate of the rate of mtDNA evolution from Kemp et al. 2007), this clade has an estimated age of 7,353 years before present (bp); however, the small number of samples produces a rather large margin of error, so that a 95 percent confidence interval (CI) encompasses a period of nearly 10,000 years, from 13,233 to 3,333 years ago. The predicted age for this clade around 7,000 years ago does not, however, seem unrealistic because a recent study of samples from a 5,000-year-old site on Santa Cruz Island demonstrated the presence of the np16093 marker among individuals belonging to haplogroup A (Monroe et al. 2010). Although using genetic data to date lineages can lead to interesting results, it does not come without many assumptions and a high degree of associated error. Continued aDNA research will continue to improve this approach, because the oldest radiocarbon-dated sample that yields a specific hap-lotype places a constraint on how quickly mtDNA evolves. To state the obvious, the temporal origin of haplogroup A with the np16093 marker detected in ancient remains from Santa Cruz Island cannot be younger than the remains in which it was detected, although it is certainly older.

Mitochondrial haplogroup B exhibits several clades that branch from the central, basal haplotype (Figure 4.4). One, defined by T→C transi-tion at np 16311, was found only among peoples speaking languages within the Numic Branch of Uto-Aztecan. A second clade, identified by a rare C→A transversion at np 16184, appeared to be present only among Yokuts peoples and their immediate neighbors. Johnson and Lorenz (2006) proposed that this clade may have arisen within ances-tral Yokutsan groups and represented a population expansion of that group within California. More recently, Schroeder et al. (2011) have discovered that the np16184 marker is also present among additional Eastern Miwok groups, as well as from Early Period (Windmiller pat-tern) and Middle Period burials in Central California (CA-SJO-112 and CA-AMA-56, respectively). These authors hypothesize that the origin of the np16184 clade may be ancestral to the Yokutsan-Utian split within the Penutian stock, which Golla (2007) has estimated to have occurred around 4,500 years bp. They used different proposed rates of mutation to calculate the age of this clade and conclude that the age of this clade likely falls between 3,000 and 5,000 years bp. Its presence at the Cecil Site (CA-SJO-112) indicates that this mutation antedates 3,000 years bp

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and supports the hypothesis that the Windmiller pattern represented an ancient Penutian-affiliated group in central California (Moratto 1984). If Yokutsan and Utian peoples entered California’s Central Valley during the Early Period, their migration may have been linked to the Altithermal, an extended period of warming that occurred during the Middle Holocene, which had a severe effect on lacustrine and other resources in the Great Basin (Sutton et al. 2007; West et al. 2007).

Another prominent clade within haplogroup B is characterized by a C→T transition at np16111 and is commonly occurring among Takic groups and their Ipai (Northern Diegueño) neighbors in south-ern California. np16111 is considered to be a “hot spot” in the mito-chondrial genome, and indeed an inspection of Figure 4.4 demonstrates that it has mutated more than once within Uto-Aztecan lineages within haplogroup B, causing reticulation in the network. It is likely that the Tubatulabal sample (arbitrarily numbered B11 in Figure 4.4) represents a separate mutation within a different clade that also includes Cahuilla sample B08. The remaining samples with the np16111 marker are likely to correlate with a G→A transition at np16483 in the second hyper-variable mtDNA region, referred to as subhaplogroup B2a (Achilli et al. 2008). Kemp and colleagues (2010) have found these two markers to co-occur among populations in the North American Southwest. They hypothesized that the np16483 marker arose among some of the first peoples to adopt an agricultural mode of subsistence in the ancient Southwest, and the demographic expansion that resulted from this econ-omy resulted in an in situ population expansion which cross-cut all lan-guage families in the region (Kemp et al. 2010; see also Bellwood and Renfrew 2002; Diamond and Bellwood 2003). Thus, the presence of np16111, which strongly correlates with np16483, among Takic groups and their neighbors may reflect ancient Takic interaction with multi-ple Southwestern linguistic groups presumably through intermarriage and admixture along the Colorado River region. The age of B2a in the North American Southwest was calculated to be 2,105 years bp with a 95 percent CI between 1,273 and 3,773 years bp (Kemp et al. 2010), thus giving us the earliest possible dates that admixture/intermarriage could have occurred with Native Californians.

conclUsion

The accumulating genetic record of California’s native peoples is provid-ing a new means to explore the past. Comparative haplogroup distri-butions show different patterns for different regions of California and demonstrate continuity between some ancient and living groups while also suggesting the replacement or coalescing of original inhabitants

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with incoming migrants. Specific mtDNA clades have been shown to correlate with certain ethnolinguistic groupings, shedding light on the origins of the diverse peoples that inhabited the California region at the time of European contact. Although the findings reported here are des-tined to be eclipsed as additional studies are undertaken, it is evident that DNA research can contribute enormously to understanding patterns of migration and genetic affiliation in California prehistory.

acknowledgMents

The authors would like to express their gratitude to the California Indian descendants who have participated in our genetic studies from 1992 to the present; to Fred Schaeffer and Kevin Hunt for assistance with assem-bling C-14 dates used in Table 4.2; and to Gary Breschini, Kari Schroeder, Sylvere Valentin, Tracey Pierre, Mark Stoneking, and Eske Willerslev for contributing information regarding mtDNA results summarized in Tables 4.1 and 4.2. Finally, we thank David Glenn Smith for his contributions and continued insights that have informed our research over the years.

note

1. PCoA is a technique used to visualize information when exploring similarities or dis-similarities in data. The genetic distance used was FST.

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