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UC Merced Journal of and Great Basin Anthropology

Title Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages

Permalink https://escholarship.org/uc/item/6g135010

Journal Journal of California and Great Basin Anthropology, 26(1)

ISSN 0191-3557

Authors Johnson, John R Lorenz, Joseph G

Publication Date 2006

Peer reviewed

eScholarship.org Powered by the California Digital Library University of California Journal of California and Great Basin Anttiropology | Vol, 26, No, 1 (2006) | pp 33-64

Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages

JOHN R. JOHNSON Department of Anthropology, Santa Barbara Museum of Natural History 2559 Puesta del Sol, Santa Barbara, CA 93105

JOSEPH G. LORENZ Laboratory of Molecular Biology, Coriell Institute for Medical Research 403 Haddon Avenue, Camden, NJ 08103

The advent of mitochondrial DNA analysis makes possible the study of past migrations among California Indians through the study of genetic similarities and differences. Four scenarios of language change correlate with observable genetic patterns: (1) initial colonization followed by gradual changes due to isolation; (2) population replacement; (3) elite dominance; and (4) intermarriage between adjacent groups. A total of 126 mtDNA samples were provided by contemporary California Indian descendants whose maternal lineages were traced back to original eighteenth and nineteenth century sociolinguistic groups using mission records and other ethnohistoric sources. In particular, those groups belonging to three language families (Chumashan, Uto-Aztecan, and Yokutsan) encompassed enough samples to make meaningful comparisons. The four predominant mtDNA found among American Indians (A, B, C, and D) were distributed differently among populations belonging to these language families in California. Examination of the distribution of particular haplotypes within each further elucidated the separate population histories of these three language families. The expansions of Yokutsan and Uto-Aztecan groups into their respective homelands are evident in the structure of genetic relationships within haplogroup diagrams. The ancient presence of Chumashan peoples in the Santa Barbara Channel region can be inferred from the presence of a number of haplotypes arrayed along a chain-like branch derived from the founding haplotype within Haplogroup A. A distinctive Haplogroup D sequence, represented by four Chumash lineages, belongs to a rare subgroup, occurring primarily among groups scattered along the Pacific coast of North and South America. This distribution is consistent with the hypothesis that an early coastal expansion occurred during the initial peopling of the .

HE HIGH DEGREE OF LINGUISTIC DIVERSITY in speaking various Uto-Aztecan languages are wedged Tnative California reflects significant migration events between Yuman societies in the San Diego-Colorado in prehistory Many of the larger, more widely-spread River-Baja California area and Chumash and Yokuts language families of North America are represented peoples in the Santa Barbara Charmel and southem San by smaU groups distributed along the Pacific Coast, as Joaquin VaUey regions. It has been presumed by nearly aU weU as by a relatively high number of compact language researchers that this "Shoshonean Wedge" was the restilt famiUes descended from more ancient migrations (GoUa of a prehistoric expansion of Uto-Aztecan peoples from 2000a, 2000b, in press). Linguist Johanna Nichols (1992) an inland region to the coast and then to the southem has described the Pacific coastal region as a "residual Channel Islands (Bright and Bright 1976; Kroeber 1953), zone," because the high ecological diversity of the region although there is considerable disagreement when this allowed for microdifferentiation and preservation of migratory event may have occurred. The most often stable linguistic communities adapted to relatively cited date is about 2,000 years ago (Moratto 1984:559); circumscribed territories. In southern CaUfomia, groups however, some researchers have suggested a much earUer

33 34 Journal of California and Great Basin Anthropology | Vol 26, No 1 (2006) period, perhaps 5,000 years ago, noting the geographic of some of these previously reported samples, and the distribution of certam distinctive items of material culture accumulation of additional samples has augmented our throughout much of the range known to have been database significantly. The expanded and refined database occupied historically by Uto-Aztecan speakers (Raab produced by oiu- continuing efforts now provides us with and Howard 2002). a clearer picture of mtDNA diversity among California The coming of age of mitochondrial DNA (mtDNA) Indian groups than has heretofore been available. genetic research by human biologists has provided an important new means of examining past migrations in many parts of North America (see Eshleman and LANGUAGE CHANGE IN Smith, in press). Mitochondrial DNA is inherited only CALIFORNIA PREHISTORY matemaUy, so over many centuries mutations graduaUy At the time of European contact, California was made accumulate in daughter lineages, making it possible to up of a veritable patchwork of local groups often caUed reconstruct ancestral types and derived lineages and tribelets (Kroeber 1963). In some regions, especiaUy in relate these to one another. AU of the mtDNA Uneages areas of higher population density supported by fishing in the Americas are associated with five recognized economies, there existed different levels of sociocultural "haplogroups" that originated earlier in prehistory integration and hierarchical relations among these among human populations in Asia (Eshleman, Malhi, local groups (Bean 1978; Johnson 1988, 2000; Jorgensen and Smith 2003; Merriwether 2002; Schurr 2004; Torroni 1980). Contributing to the cultural mosaic represented 2000). Observations of genetic affinities between ancient by different tribelets and regional differences in socio­ peoples and ethnographicaUy documented groups have political organization was a high degree of linguistic been matched with archaeological and linguistic evidence differentiation. Some 88 distinct languages were spoken to reconstruct population movements that led to pattems between the southem tip of Baja California and Oregon observed at the time of European contact (Eshleman, (Goddard 1996a, 1996b; Laylander 1997; Mithun 1999). MaUii, and Smith 2003; Eshleman et al. 2004; Kaestle and These have been classified into fourteen language famiUes Smith 2001; Kemp et al. m press; Lorenz and Smith 1996, and seven isolate languages. A number of these famiUes 1997; MaUii, SchuLz, and Smith 2001; Malhi et al. 2002, and individual languages traditionaUy have been grouped 2003, 2004; O'Rourke, Hayes, and Carlyle 2000; Rubicz into two macro-units or superfamilies, Hokan and et al. 2003; Schurr 2004). Penutian, although the reaUty of these larger taxonomic Over the past thirteen years, the two coauthors entities remains hypothetical and is a subject of current of this study have collaborated to study the mtDNA investigation and debate among specialists in comparative from 126 separate California Indian matrilines (direct linguistics. One interpretation is that these macro-units female lineages). These lineages have been traced appear to represent ancient language families that through genealogical methods to specific native groups, became dispersed and evolved into descendant famiUes mostly in south-central and southern California, thus and isolated languages through Unguistic processes (GoUa providing a new database that elucidates the mosaic of 2000a). Linguists and archaeologists have been interested genetic relationships in the region. This paper will use in explaining the distribution of languages in the Pacific the sequences of the principal mtDNA hypervariable Coast region and adjacent Great Basin and have devoted segment (HVSl) of 121 of these lineages in its analysis. much effort to reconstructing linguistic prehistory and EarUer pubUcations of portions of these data (Lorenz and proposing likely cultural processes (e.g., Bettinger and Smith 1996,1997) and subsequent unpubUshed additions Baumhoff 1982; Breschini 1983; Bright and Bright 1976; have been used in a number of comparative studies GoUa 2000b; HiU 2002; Hughes 1992; Jackson 1989; King of native North American genetic relationships (e.g., 1986; Krantz 1977; Levy 1997; Madsen and Rhode 1994; Eshleman 2002; Eshleman et al. 2004; Kaestle and Smith Moratto 1984; Nichols 1988;Whistier 1977,1978,1988). 2001; Kowta 2003; Malhi et al. 2002, 2003; O'Rourke, It is often assmned that populations speaking related Hayes, and Carlyle 2000). Continuing genealogical languages might share genetic lineages in common, research has revised the ethnoUnguistic group affiUations because they once had belonged to the same group ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnson / Lorenz 35 prior to linguistic differentiation. There are a number of 1. Ancient populations that retain their residence factors, however, that make the correlation of languages in one region over many miUeimia wiU exhibit and less than straightforward group-specific markers and branching, chain- (Moore 1994; Sims-WUUams 1998). Although m general Uke pattems of mitochondrial DNA variation language famiUes often do share similar genetic Uneages with all descendant lineages preserved in populations around the world (CavalU-Sforza 2000; within the group. Populations that bud off Cavalli-Sforza, Menozzi, and Piazza 1994), there are and migrate elsewhere usually only retain plenty of exceptions to this rule because language a subset of the range of variation present in ttansmission can occur for reasons having nothing to do the ancestral population. In addition, ancient with the physical movement of large numbers of people groups in California may be linked to some (e.g., Nasidze and Stoneking 2001). of the first expansions that peopled the An extensive literature has developed in recent Americas if they are found to possess rare years regarding models of language spread and replace­ lineages held in common with other early ment (Bakker 2000; BeUwood and Renfrew 2002; Dbcon migratory groups. Such rare, ancient lineages 1997; GoUa 2000b; Foley 2004; Jones 2003; Nettle 1999, would not necessarily be present among later 2000; Nichols 1992, 2000; Renfrew 1987,1992, 2000). peoples who entered California subsequent Based on our review of this Uterature, we propose that to the original founding populations. the foUowing four general scenarios are Ukely to account 2. Population replacement will result in for most of the linguistic changes that occurred in differences between mtDNA Uneages found prehistoric California: among peoples Uving in the region today and 1. Initial colonization by Paleouidians foUowed samples from prehistoric burials of earlier by gradual changes through time due to periods. A spreading population also produces isolation or differentiation along dialectal a "star-like" pattern of descendant lineages continua. from the ancestral type, resulting from genetic drift and isolation as descendant subsets of 2. Population replacement through immigra­ the group move into geographicaUy separated tion, with one group forcing another to areas. In cases of population replacement, abandon its territory, because of greater remnant lineages may exist here and there, numbers, technological advances, more representing the earUer group, just like certain effective subsistence strategies, or some lexical, phonological, or grammatical features aspects of sociopolitical organization that may become incorporated mto the language give the incoming group an advantage over of the incoming group. the former inhabitants. 3. Elite dominance will result in just the 3. Elite dominance, whereby an incoming opposite genetic pattern from that occurring group estabUshes hegemony over the original with population replacement. In cases of inhabitants, without displacmg them, initiating eUte dominance, the genetic pattems wiU not an adoption of the language spoken by the change significantly from those found among new poUtical leadership. prehistoric peoples in the region, although 4. Intermarriage of adjacent groups over language change took place. As was the an extended period of time, leading to case in number 2 above, a certain Unguistic UnguisticaUy mixed conununities that would substrate may survive of the earUer language shift from one language to another over spoken in the region. several generations. 4. Intermarriage of adjacent groups wiU result Furthermore, we expect each of these foiu- scenarios in shared genetic Uneages, even though the to have resulted in different genetic pattems: peoples themselves speak imrelated languages. 36 Journal of California and Great Basin Anthropology | Vol, 26, No, 1 (2006)

Depending on the origins of the respective Golla (in press) has summarized our current groups prior to their settlement adjacent to understanding of California's linguistic prehistory— one another, one might be able to detect following more than a century of investigation—to differences between prehistoric pattems in reconstruct probable scenarios that resulted in the the region and detemune which Uneages were configuration of languages existing at the time of European once associated with each group prior to their contact. Some of these proposed reconstructions are intermarriage with one another. amenable to testing using mitochondrial DNA research. The sample utUized for this study (see below) is most Based on these different expected outcomes, one can informative for Central and Southem California, because examine the actual pattems of mtDNA variation among we have not yet had the opportunity to conduct research CaUfomia ethnoUnguistic groups in order to reconstruct among Northem California's indigenous peoples. Only six past processes of cultural change and compare these to lineages characterized here pertain to the region north of aspects of the archaeological record. It is important to pomt San Francisco (see "Sample Descriptions" below). Thus, out, however, that informative as they potentiaUy can be, GoUa's hypotheses pertaining to the Algic, Athabaskan, mtDNA pattems present among CaUfomia Indians were Wmtuan, Maiduan, and Yukian famiUes and the dispersed very dependent on the form of post-marital residence Northem Hokan languages are not amenable to testing practiced by the different groups. Since mtDNA is only using oiu- dataset. inherited maternally, groups that practiced patrilocal Three groups in the Central and Southern post-marital residence would be Ukely to share matemal California regions, however, probably do possess large lineages with adjacent groups, whereas groups in which enough samples to begin to inform us about past genetic matrUocal post-marital residence predominated would relationships among ethnoUnguistic groups. These preserve mtDNA distinctiveness over many generations. mclude the Chumashan family, the Uto-Aztecan family, Our abiUty to differentiate between the different cultural and the hypothesized Yok-Utian branch of the Penutian processes that led to language change and distinctive superfamily. Also, a modest number of samples from mtDNA pattems would be obscured in patrilocal cases central and southem Hokan peoples (SaUnan, Esselen, where wives moved across linguistic boundaries to take and Yiunan-Cochirm') permit comparisons with adjacent up residence in their husband's local group. groups that possess larger numbers of samples. Golla GeneraUy, two pattems of social organization prevailed has proposed that because of its linguistic distinctiveness in Native CaUfomia: (1) group affiUation based on bilateral and lack of estabUshed relationships to other language descent in Northem CaUfomia, and (2) group affiUation fanulies of the Americas, the Chumashan family might based on patrilineal descent with concomitant patrilocal weU constitute one of the "basement" language famiUes residence m Central and Southem CaUfomia (Jorgensen of CaUfomia (GoUa 2000c). In contrast to Chmnashan 1980; Kunkel 1976). It has been recently argued that even peoples, the spread of Uto-Aztecan languages into in Northem CaUfomia, the predominant form of social CaUfomia appears to have occurred during a later period organization was "patrifocal," although greater flexibiUty of prehistory, perhaps beginning four mUlennia or more in post-marital residence occiured there (Burton, Moore, before present. One currentiy accepted scenario, based on and Ronmey n.d.).The principal exceptions to the overaU linguistic evidence, derives their origins from a region in patrifocal emphasis in aboriginal California societies Mexico. The resultant dispersal of Uto-Aztecan languages were the Central and Island Chumash peoples, where a northward occurred with the spread of maize agriculture matrilocal residence pattem predominated (Johnson 1988; into the American Southwest. As populations budded 2001). Given these differences in CaUfomia Indian post- off from these agricultural commimities into desert areas marital residence preferences, one would expect a greater unfavorable for growing crops, the descendant groups degree of mitochondrial DNA distinctiveness among returned to a himting and gathering adaptation, spreading Chumashan groups than might prevail elsewhere in into the southem San Joaquin VaUey and adjacent southem Native California, where patrilocal residence or bUateral Sierra Nevada (Diamond and BeUwood 2003; HiU 2001, kin groups occurred. 2002). Both SaUnan and Northem Chumash languages. ARTICLE { Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Jotinson / Lorenz 37 as weU as those of some other Central CaUfomia groups, 3. Yokutsan and Uto-Aztecan groups, being show evidence of prehistoric lexical borrowings from more "recent" arrivals into California, wiU "Old California" Uto-Aztecan languages prior to the share mtDNA lineages with other regions differentiation and expansion of the Numic subfamily of origin as well as less common mtDNA (Klar 1980; Nichols 1988;Tumer 1987).The later movement lineages from groups absorbed during the into Southem CaUfomia of Uto-Aztecan peoples has been process of expansion. proposed to have occiured about two miUeimia ago, and to 4. Intermarriage between adjacent groups have originated from a region in the vidnity of the southem from different language famiUes, especiaUy San Joaquin VaUey (GoUa, in press; Moratto 1984). those practicing patrilocal post-marital Within the hypothesized Yok-Utian branch of the residence, wiU result in a certain amount of Penutian macro-unit, GoUa has proposed two migrations: sharing of mtDNA Uneages. Such adjacent one that introduced Rroto-Utian into the Sacramento- groups that exhibit the highest degree of San Joaquin DeUa region some four miUeimia ago, and linguistic influences, through features such a second migration within the Late Period that brought as phonological convergence or lexical Yokutsan into the Central Valley. He proposes that borrowings, wUl tend to be indistinguishable the ancestral homeland of both these families and of from a standpoint of mitochondrial DNA the Penutian superfamily generally was likely to have Uneages, reflecting intermarriage. been in the Plateau region and in portions of the Great Basin. This idea was tested in part by Kaestle and Smith (2001) in their research contrasting ancient Great Basin population samples with samples obtained from modem SAMPLE DESCRIPTIONS Numic peoples, using a subset of the data collected The research design for this study is unique among for this study for their characterization of Califomian Native American mtDNA studies conducted to date groups. Yokutsan languages show substratal influences in that extensive genealogical docmnentation has been from SaUnan (Golla, in press), suggesting that some undertaken using ethnohistorical records. This approach mitochondrial lineages from an absorbed earUer group was necessary because the relocation of many California may be present in Yokuts populations. groups to the missions during the Spanish colonial period A number of predictions may be derived from this and the subsequent demographic decUne caused by the discussion pertaining to language changes in aboriginal introduced European diseases led to intermarriage and CaUfomia: coalescence of what had been independent tribal entities. Present-day descendants of California Indians may 1. Chumashan populations will be genetically identify themselves with a particular tribal designation; distinctive compared to neighboring groups, however, genealogical research may reveal that the because of the likelihood of their ancient matriline of these individuals descends originaUy from a presence in the Santa Barbara region and woman from a different native group who had married a their matrilocal residence pattern, which man from the person's current tribe. If the anthropologist would tend to preserve mitochondrial DNA did not undertake genealogical research, many of the lineages within the region. mtDNA samples would be misassigned to incorrect 2. Rresiuning that the distribution of languages ethnoUnguistic units, thus obscuring original patterns within the Hokan and Penutian superfamiUes of genetic differences and similarities that had once represent ancient population spreads existed among California's native groups. By using in CaUfornia, the descendant populations mission registers. Bureau of Indian Affairs (BIA) records, speaking different languages found within ethnographers' notes, censuses, and other sources, each of these superfamilies will harbor a relatively high degree of precision is obtained in ancestral mtDNA Uneages, even though they determining the origin of most of the Uneages sampled. are widely separated geographicaUy.' It should be emphasized, however, that in the end our 38 Journal of California and Great Basin Anthropology | Vol, 26. No, 1 (2006) efforts have only achieved a small sampling of the In certain cases, especially for Chumash and diversity that must once have been present m aboriginal other Central CaUfornia Coast lineages, the research California. Inevitably, the demographic collapse of was conducted in reverse; that is, mitochondrial DNA the nineteenth century resulted in the loss of many lineages could be traced from known ancestors Usted in mitochondrial DNA lineages. Although a relatively high mission records to famiUes Uving in the region today. These degree of confidence may be placed in oiu^ affiUation of individuals were then approached about their participation specific mtDNA Uneages with particular tribal groups, in the study. So rather than working backwards through these may not be tmly representative ui cases where only the records, these people were contacted because we had a few lineages survive. followed their lineages forward through documentary The first samples obtained for this study were research from mission times to the present. In sum, mission collected primarily among Chumash, SaUnan, Yokuts, register research produced a total of 46 mitochondrial DNA and Kitanemuk descendants in 1992. Sampling and Uneages in which the town or viUage could be determined documentation of lineages has been ongoing since that where someone's female ancestor had been bom more date. From 1992-1998, hair foUicle samples were the than two centuries previously. It should be emphasized that principal means employed to obtain the mitochondrial our study differs from most other studies reported in the DNA. Beginning in 1998, buccal swabs became the Uterature in that we are sampling lineages that have been preferred method. AU samples were stored in a freezer determined to be derived from independent matrilines before being processed in the lab. In addition to California as far back as can be traced through historical records. Indian samples collected, an even greater number of Other studies have used samples from tribal populations mtDNA samples from people of Mexican American and without extensive knowledge of genealogies, from medical early colonial Spanish-Mexican famiUes were obtained cUnics where degrees of relatedness were tmdociraiented, and wiU be discussed in a separate pubUcation.^ or from prehistoric cemeteries of unknown familial Initial interviews with donors were conducted background. Thus measures of frequency or diversity regarding knowledge of their ancestry, with particular calculated from these different sampUng strategies are attention to their female lineage. Based on this not directly comparable to each other or to the findmgs information, genealogical research was conducted usmg reported here. various ethnohistoric sources. UsuaUy a person was able Table 1 classifies the 126 mitochondrial DNA lineages to supply information on the name and place of birth sampled in this study according to Unguistic groupings, of their mother and maternal grandmother, often with and Figure 1 shows the geographic distribution of additional details on those individuals' dates of birth, these samples. Inevitably, not all of the ethnoUnguistic marriage, and death. Because California Indian famiUes affiliations could be determined precisely for aU of the have had to supply genealogical data to the BIA for female ancestors of the matrilines that gave rise to these enroUment piuposes in the twentieth century, begiiming samples. In particular, the ethnoUnguistic affiliations with the CaUfomia Indian Jurisdictional Act of 1928, they of eleven lineages could not be determined with a often can provide information about earUer generations high degree of confidence, even though in some cases as well. Genealogical data supplied by families were these descended from women mentioned in records checked against information provided on the original from particular missions. In these instances, usuaUy the applications for the 1928-1933 enrollment and/or woman's name could not be matched with certainty county birth and death records, church baptismal and to a particular baptismal entry, so her forebears were marriage records, census records, and so forth. The goal tmknown. In other cases, the women were described to in genealogical research was to link California Indian be of CaUfornia Indian ancestry in BIA records of the descendants of today with their ancestors baptized at 1928-1933 eiu-oUment and/or ethnographic records, but the different missions. The mission records were then tribal affiUations were unmentioned. FinaUy, there were searched to determine insofar as possible the original a few cases of Indian chUdren being raised in non-Indian native rancheria (town or viUage) where the earliest households, so their parents' names and tribal affiUations female ancestor in the direct matriline had been bom. were tmknown. If there was no documentation at aU to ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnson / Lorenz 39 substantiate a person's claim of possessmg a CaUfomia Table 1 Indian genetic inheritance through a direct female Uneage, such samples were omitted entirely from this analysis. In DISTRIBUTION OF CALIFORNIA INDIAN MITOCHONDRIAL DNA SAMPLES ACCORDING TO RECOGNIZED LINGUISTIC DIVISIONS some of these last cases, it is expected that such evidence may eventuaUy be discovered, but to avoid clouding the Number of Documented Number of Haplotypes Not results, it was deemed better to proceed wdth the analysis Linguistic Group Lioeages Haplotypes Unique to Group Sequenced of only those samples where Utile doubt existed. Ciiumaslian Family For the most part, the samples in this study were Northern (Obispeho) 3 1 0 1 categorized according to the most recent linguistic Central classification of Goddard (1996a, 1996b) and Mithun Purlslmerio 2 2 0 0 (1999), with some minor revisions based on the Ineseiio 5 3 0 G Barbareiio 4 4 2 0 classification used by GoUa (m press). Except for Uto- Ventureiio 6 6 5 0 Aztecan and Chumashan, sample sizes were relatively Island (Cruzeiio) 1 1 0 0 Chumashan Totals 21 13 10 1 smaU for most language famiUes, so language families within the Hokan and Penutian superfamilies were Hokan Supetlamiiy (Proposed) grouped together within these macro-imits for piuposes Achumawi/Atsugewi 1 1 0 0 Esselen 1 1 0 0 of comparative analysis. It has long been assumed that Salinan 6 5 3 0 the Hokan and Penutian superfamilies represented Yuman-Cochimi Family population spreads at separate times ui prehistory, and Ipal 7 6 4 0 Tipai 1 1 0 0 our working hypothesis was that distinguishing genetic Yuma 1 1 1 0 traces of these ancient migrations might be possible. Cochimi 1 1 0 0 Hokan Totals 18 15 8 0 Grouped according to maximal Unguistic relatedness, our mtDNA dataset is represented by four principal divisions Penutian Supetlamiiy (Proposed)1 (Chiunashan, Hokan, Penutian, and Uto-Aztecan), with Wintuan Family Wintu 2 2 1 0 single samples obtained from each of two groups along Southern Patwin 1 1 0 0 the northem coast (Yiu^ok and Coos). Because the quaUty Utian Family of this genetic record is dependent on the reUabiUty of the Sierra Miwok 4 4 1 0 Southern Costanoan 4 4 3 0 ethnic affiUation of each of the samples, we have mcluded Yokutsan Family descriptive information on the particular Uneages sampled Nim Yokuts 13 8 5 1 Buena Vista Yokuts 3 2 0 0 within each group. Penutian Totals 27 17 10 1

Origins of the Chumash Samples Uto-Aztecan Family Tubatulabal 4 4 2 0 The Chumashan family is composed of at least six Numic Branch languages in three branches: Northem (Obispeiio), Central Kawaiisu 5 3 1 0 Mono 4 4 3 0 (Rurisimeiio, Ineseiio, Barbareiio, and Ventureno), and Takic Branch Island (Cruzeiio) (Klar, Whistler, and McLendon 1999). Gabrielino 2 2 0 0 This fanuly was formerly included in the Hokan macro- Kitanemuk 3 3 1 0 Serrano/Vanyume 4 3 2 0 unit (Heizer and Elsasser 1980; Kroeber 1953), but it has Cahuilla 5 5 3 G since been determined to constitute an isolated language Cupeno 2 2 0 G family with an ancient presence in California (Mithun Luiseiio 18 9 4 2 Uto-Aztecan Totals 47 25 13 2 1999; also see GoUa, in press). Samples from 21 mtDNA lineages, representing the territories of aU six Chumashan Otiier Pacific Region Groups languages, were obtained for this study. Nineteen of these Coos 1 1 1 G Yurok 1 1 0 0 could be documented as descendmg from women who Uncertain Tribal Ancestry 11 9 5 1 resided in specific native towns and viUages (Fig. 2). These Other Groups Totals 13 11 6 1 comprise aU but one of the known surviving Chumashan Total California Indian Sample 126 67 40 Journal of California and Great Basin Anthropology { Vol. 26, No 1 (2006)

Figure L Map showing linguistic afiiliations and distribution of samples obtained for mtDNA analysis. mitochondrial DNA lineages.^ Five additional lineages samples derived from these five lineages have been were traceable to forebears Uving in the vicinities of the omitted from the Chumashan totals and included in the Chiunash missions in the mid-nineteenth century; however, "Uncertain Tribal Ancestry" category in Table 1 and our attempts to trace the tribal origin of these matrilines has subsequent analyses. not yet been successful. One carmot be certain whether The samples derived from the 21 documented these Uneages originated among Chimiash populations or Chumash matrilmes include three Northern Chumash from Yokuts or other CaUfomia Indian women who had lineages, two Rurisimeiio Uneages, fiveIneseii o Uneages, moved into the area from elsewhere. Thus, the mtDNA four Barbareiio lineages, six Ventureiio lineages, and ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnson / Lorenz 41

a'.."*!: LMI Obispeno^"*" '^^v^ Salinan /

,*, Tipud) \^ /

Mfssion • w S,.Btte^aventura ^ Mupu(1) -^ o ^ \ (1) '^ Shisholop 'Y», Kayiw.sh(1)» jm„^„l N • Towns witti mtDNA lineages V • IS, Fernando Ba rbareno (1) \ • • Ottier Villages \^uS) • . j'GabTrieiino (2) Number of mtDNA lineages . Hichtmtn • • . from town • • ... •() • ".. *

0 r u z e n o 0 5 10 20 30 40 I Kilometers

Figure 2. Origins of mtDNA lineages from Chumash towns and villages. one Cruzeiio lineage. The Northern Chumash samples from women bom at Kashtayit (Santa Anita Creek) and include one from a woman born in Tipu in the upper 'Onomyo (Gaviota) along the western Santa Barbara Salinas River watershed, one from a woman from Chaimel coast. The five Ineseiio matriUnes descend from Tsipxatu at AvUa Beach, and one from a woman Usted as two women born at Kalawashaq', one woman from being from Mission San Luis Obispo in the appUcations Soxtonokmu', one woman from Mi'asap, and one woman of her descendants in the 1928-1933 CaUfomia Indian from Shnaxalyiwi. The four Barbareiio lineages descend enrollment. Potentially, this last mentioned individual frora two women from Mikiw (Dos Pueblos), one woman could also have been a Yokuts woman, because many from Syuxtun (Santa Barbara), and one woman from people from Valley Yokuts tribes were baptized at Shuku (Rincon). Five Ventureiio samples descend from San Luis Obispo; however, her mention among John women bom at Shisholop (Ventma), Mat'ilha (MatUija), Harrington's Obispeiio field notes has led us to include Mupu (Santa Paula), Kayiwish (Cayegues), and Muwu her among the Northern Chumash Uneages until such (Mugu). A sixth Ventureiio lineage descends from a time as her identity in mission records can be estabUshed woman named Man'aTarango, who was reported by John with certainty. Harrington to have spoken the Ventureiio Chumash AU but one of the Central Chumash samples and language, but whose identity in the mission records has the single Cmzeiio sample are traceable through their proven elusive. The Cruzeiio matriline stems from a direct female lineages back to their original ancestral woman bom at Hichimin on Santa Rosa Island who was towns and viUages. The two Rurisuneiio Uneages descend baptized at Mission Santa Ines. 42 Journal of California and Great Basin Anthropology | Vol. 26, No. 1 (2006)

Origins of the Hokan Samples potential Salinan mitochondrial DNA Uneages have been With two exceptions, aU of the samples pertaining to the detemuned to exist today. Hokan superfamily were from speakers of the SaUnan Yuman-Cochimi Samples. Seven of the ten samples language of Central California (six samples) and the gathered from the descendants of peoples speaking Yimian-Cochimi fanuly of Southem CaUfomia and Baja Yuman-Cochimi languages came from the Ipai (Northem California (ten saraples). These two Hokan language Diegueno), a group that often intermarried with its areas were separated by Chumashan and Uto-Aztecan Takic (Luiseno and Cupeiio) neighbors. Because of peoples at the time of Eiu^opean contact. One sample was this intermarriage and because these groups practiced obtained from an mdividual who descended through her patrilocal post-marital residence, it was to be expected direct female Une from a woman baptized at Mission San that some mitochondrial DNA haplotypes would be Carlos from the Excelen tribelet, from which the name shared between Yuman and Takic peoples, reflecting the Esselen was derived. The single sample from a Northem movement of women across Unguistic boimdaries.Two of Hokan lineage was obtained from a descendant of a the Ipai samples were from people who belong to Luiseiio woman who was from the Pit River area, with Utile else groups today. Their direct female Unes, however, traced known regarding her background. In the absence of soUd back through the Mission San Luis Rey records to women genealogical evidence, this Uneage has been presiuned to from the Ipai ranchen'as of Bataquitos and Tahui (Tawi) be either Achumawi or Atsugewi in origin, with the caveat who had married Luiseiio husbands at the mission. The that it could also be descended from a woman who had remaining five Ipai samples came from people descended moved into the Pit River area from a neighboring group. from the nineteenth century reservation communities Salinan Samples. Samples from six Salinan lineages of Santa Ysabel, Mesa Grande, and San Rascual. The were obtained for this study. In addition to these, two single Tipai (Southern Diegueiio) sample was from an samples from people whose ancestors were baptized individual whose direct female line was traceable back at Mission San Antonio appear to have come from to a woman from Mission San Miguel in northem Baja Northem VaUey Yokuts women, despite having survived California. The Yuma sample was from a woman who among people who today consider themselves SaUnan had moved from southem Arizona sometime prior to her or Northem Chumash. These two samples are described marriage at Mission San Diego. Her descendants later below in the Yokuts section. Using California Indian intermarried with the Luiseiios.^ The Cochimi sample enrollment records and/or mission records, four of the was obtained from an individual of California Spanish six SaUnan lineages could be traced to ancestral viUages descent whose direct female Uneage descended from a where the direct female ancestor bad been bom. These neophyte woman at Mission Santa Gertrudis in Baja were: Lima in the upper San Antonio valley, Isley in CaUfomia in the latter part of the eighteenth century. the coastal district of Lamaca, Monet along the upper Nacimiento River, and Sicpats in the northern Carrizo Origins of the Penutian Samples Plain.'* The last mentioned sample could potentiaUy be About 60 percent of our 29 samples within the from a Northem Chumash lineage.^ The two remaining Penutian stock derive from ancestors who spoke Yokutsan lineages descend from Salinan families whose direct languages; however, a few samples pertain to the Wintuan female lines could be traced back to neofitas at Missions family and some were Miwok-Costanoan in origin. The San Antonio and San Miguel who were married in the specifics regarding these Uneages are provided below. decades following the secularization of the missions. Wintuan Samples. The Wintuan family consists of Unfortunately, the baptismal records for neither woman Wintu, Nomlaki, and various Patwin languages. Two could be certainly identified. Either or both of these two Wintu samples, one Namtipom and the other Wirmemum, matrilines could therefore potentially be of Northern were documented using the California Indian enrollment Yokuts origin, because of the large number of people records. The Southem Patwin sample was obtained from from the San Joaquin Valley who were recruited to an individual whose female ancestor was baptized at these missions after the Salinan population had been Mission San Jose from the original Napa tribelet. This proselytized. Other than these six matrilines, no other woman subsequently married a Costanoan man, and ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnson / Lorenz 43 their descendants today are affiUated with an "Ohlone" Livingston), Zucuy (unlocated), Chauyat (unlocated), (Costanoan) group. Chawchila, and Chukchansi.^ Two other Northern Miwok-Costanoan (Utian) Samples. Four Miwok VaUey lineages, one at San Juan Bautista and one at San and four Costanoan lineages have been sampled for this Antonio, could not be traced to named groups, reflecting study. The Miwok samples are aU from the interior region, the tendency for some missionaries to designate Yokuts apportioned as foUows: one Northem Sierra Miwok, one individuals generaUy as "Tulareiios," without regard to Central Sierra Miwok, and two Southem Sierra Miwok. their specific origin. The Southern VaUey and Foothill The four Costanoan samples are aU from southem groups. lineages were descended from women from Nutunutu, These include two lineages descending from women Tachi (3), Yawdanchi, and Yawelmani (2) groups. The baptized at Mission San Carlos (one Ensen, one Tucutnut) donors of most of the Southern Valley and Foothill and two lineages descending from women mentioned in Yokuts samples are members of federally recognized the Mission San Juan Bautista records. One of the San Yokuts tribes. Juan Bautista samples descended from a woman born m the Mutsun tribelet of Raigssin (Raicines); the other Origins of the Uto-Aztecan Samples has not yet been determined. The last mentioned sample The largest number of samples collected for this study could conceivably be of Yokuts origin instead of Mutsun, originated among groups speaking Uto-Aztecan because the female ancestor of this Uneage has not been languages. Linguists classify California's Uto-Aztecan identified with absolute certainty in the mission records. languages into three major branches: (a) Tubatulabal, Yokuts Samples. Whistler and GoUa (1983) have a single language; (b) Numic; and (c) Takic (Goddard divided the Yokutsan fanuly into three principal branches: 1996b; Mithun 1999:539). Most of our samples (33) Poso Creek (Ralewyami), Buena Vista (Tulamni and descend from women who spoke languages in the Takic HometwoU), and Nim Yokuts (FoothiU and VaUey tribes). group, but samples were also obtained from people Our samples do not include Ralewyami lineages, but whose female ancestors spoke Tubatulabal (4) and two of otherwise they are fairly weU distributed among groups the Numic languages (9). The Takic languages in southem located throughout the San Joaquin VaUey region. People California are regarded as having been introduced from the various Yokuts tribes were baptized at many relatively late in prehistory. California Missions, from San Jose in the north to San Tubatulabal and Numic Samples. Tubatulabal and Femando to the south. Six of our Yokuts samples could be several Numic languages are spoken by neighboring traced back to ancestors baptized at the missions (two at groups in the southem Sierra Nevada region. There has San Juan Bautista, three at San Antonio, and one at Santa been considerable intermarriage among these groups, Ines). The remaining Yokuts samples were obtained from who practiced patrilocal post-marital residence, so that people whose lineages could be documented through the mitochondrial DNA lineages would be expected to be California Indian enrollment records and genealogical shared among them to a certain extent. Moreover, it is data coUected by various ethnographers (e.g., Gayton- difficult to trace female luieages back beyond the mid Spier n.d.; Harrington 1985). to late nineteenth century, because the native peoples The Buena Vista Yokuts samples came from one of this region were generaUy not drawn into the mission Tulamni and two Hometwoli (Taneshach) Uneages. system and therefore names and viUages of origin were One of the HometwoU lineages survives today among not recorded for earlier generations. CaUfornia Indian Chumash descendants, resulting from a marriage between enrollment records, genealogical data in ethnographic a Taneshach woman and an Ineseiio man. The two other fieldnotes (e.g., Harrington 1985,1986), the Kelsey census lineages from the Buena Vista group survive among of non-reservation Indians in 1905 (Kelsey 1971), and U.S. Yokuts descendants who settled at the Tejon Indian census records provided the principal means of tracing community. The Nim Yokuts samples were relatively female lineages. evenly divided between northem and southern groups. Four Uneages were determined to be Tubatulabal Five Northem VaUey Uneages could be traced to women in origin: three directly traceable to women born in from particular groups or ranchen'as: Silelamne (near the upper Kem River watershed, and one reported as 44 Journal of California and Great Basin Anthropology | Vol. 26, No. 1 (2006)

Bankalachi from Tulare County. Nine Numic samples Cupa.The second descended from a weU-known Cupeiio were coUected in the course of our investigation. Three family documented by Strong (1929:194). of these were Westem Mono (Monache), one was Mono The largest number of samples obtained from from the eastern side of the Sierra Nevada (Owens VaUey descendants of speakers of any single language was Paiute), and five were Kawaiisu. Two of the Kawaiisu seventeen obtained from Luiseiio Indians. Because samples descended from women from Kelso VaUey, one the original baptismal, marriage, and burial records for was from a woman from the Paiute Mountain area, and Mission San Luis Rey have been lost for more than two were from women from the vicinity of Tehachapi. a century and a half, a combination of ethnohistoric Takic Samples. Takic languages are divided into sources was used to trace the lineages of those who two subgroups: Serran (Serrano-Vanyume, Kitanemuk, provided samples. The most important documents for Tataviam, Gabrielino) and Cupan (CahuiUa, Cupeiio, providing data on Luiseiio genealogies were the two Luiseiio). Our samples included 9 lineages from Serran surviving padrones (census books) of Mission San Luis groups and 28 lineages from Cupan peoples. The three Rey, the 1852 CaUfomia State Census, nineteenth century Kitanemuk samples came from families belonging to parish books, BIA heirship records, and California the Tejon Indian community. Two Vanyume lineages Indian enroUment records (Johnson and Crawford 1999; descended from women baptized at Mission San Femando Johnson and O'NeU 2001). Using these various sources, from the ranchen'a of Topipabit, located in the VictorviUe ten matrilines could be traced to source ranchen'as Usted Narrows. The two Serrano lineages came from women in the padrones, five could be traced to late nineteenth who were affiliated with Morongo Reservation near century reservation communities, and two were of Banning. One of these was from a family who had come undetermined origin (Figure 3.). Most of the Luiseiio from the Mission Creek area near Morongo VaUey. The samples where duect female Unes could be traced were direct female ancestor of the remaining Serrano lineage derived from groups originaUy located in the vicinity of has not been determined with certainty and conceivably Raloraar Motmtain (three from Cuqui, two from Toulepa, could be of CahuiUa derivation instead. The two Uneages one from Temecula, one from Pimixga, and one from included in the Gabrielino group descend from women Aguanga). Only two matrilines descended from women affiliated with Mission San Gabriel and Mission San who Uved closer to the coast, both from the rancheria of Femando, respectively. The Gabrielino Uneage is traceable Topome (Topomai) on the Santa Margarita River, which through the mission records to a woman from Quinquina, was the largest of the Luiseiio poUties. San Clemente Island.^ The Femandeiio sample has not been successfully associated with a female ancestor baptized from a native rancheria, and could conceivably RESULTS AND OBSERVATIONS have descended from a Tataviam speaker.' The comparison of mtDNA haplogroup frequencies In the Cupan division, five samples are traceable to provides a low-resolution means of differentiating Cahuilla ancestors, using CaUfornia Indian enroUment between populations of dissimilar ancestral origins. For records and ethnographic information. Two matrilines most samples, restriction analysis had been conducted were traceable to female ancestors Uving on the CahuiUa prior to sequencing to determine mitochondrial Reservation near Anza, and thereby are probably from haplogroup affiliation. Much more information was the Mountain Cahuilla group; one is traceable to an obtained by analyzing sequences obtained from the ancestor at Soboba Reservation; one to an ancestor principal non-coding, hypervariable segment (HVSl) baptized at Mission San Gabriel frora the Pass CahuiUa of the mtDNA molecule. When two individuals possess ranchen'a of Reatopa (Pihatapa); and the last to a Desert identical HVSl sequences, they are said to belong to the Cahuilla woman born near Indio, who had moved to same "haplotype" and share a common ancestor within a Morongo with her Serrano husband. One of the two particular haplogroup. The techniques used in sequencing Cupeiio samples was initially thought to be Luiseno, for this study were those described by Lorenz and Smith but when traced through San Luis Rey mission records (1997) and Lorenz et al. (2005). After sequences were proved ultimately to have descended from a woman from obtained, these were checked backwards and forwards ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Jolinson / Lorenz 45

T-Enj—rrT3—r- II iM i pi Pimixga (1)

^...--'•"Tbulepa (2) Pechanga Aguanga (1) ^ f Reservation

La Jolla gaprvation Cuqui/ •^^"•™^2) (3)1 • ICupa (m>

*""M ifyj mini ' >^ ^San n rascuaPascuall \ H Santa 'i^belV / R^ervationd) ^i ReserveftiSn (2)

^ • Towns witti mtDNA lineages f :?»^i|^^J^ Jteservation (2)

• Otiier villages Bataquitos (1) (2) Number of mtDNA lineages ipai from town or reservation

0 2.5 5 10 15 ^E^H^=^^^^^^ Kilometers

Figure 3. Origins of mtDNA lineages from Luiseiio, Cupeiio, and Ipai communities. in many subsequent trials in order to be sure the results discovered among the lineages we sampled.^^ Our results were accurate. In a number of instances, more than indicate that there are clear differences between three one individual from the same matriUne had provided of the principal language famiUes present in central and mtDNA samples, and so this provided a further check southem CaUfomia in terms of haplogroup frequencies on our sequences.^" Our tables for each haplogroup (Figure 4). Haplogroups A and D predominate among report aU variable nucleotide positions (np) between the Chumash, haplogroups B and D prevail among the np 16051 and np 16362 that deviated from the Cambridge CaUfornia Renutians, and haplogroups B and C are in Reference Sequence. These variable positions thus serve the great majority among Uto-Aztecan groups. The as distinctive markers to study phylogenetic relationships CaUfomia Hokan sample, heavUy weighted by its Yuraan among various California Indian Uneages.^^ component, also has a high proportion of haplogroups B and C. The similarity in haplogroup distribution between Comparative Haplogroup Frequencies Yuman and Uto-Aztecan groups appears to be the result Table 2 summarizes the haplogroup frequencies observed of extensive intermarriage between the Ipai and Luiseno, in individual ethnoUnguistic groups with totals for the each of which comprise a pluraUty of samples within their language families and superfamiUes used for broader respective groups. The most striking contrast is between comparisons.^^ All of our samples fell within the four Chumashan and Uto-Aztecan peoples. Comparing predominanthaplogroupsfoundamongAmerican Indians just these two groups in a 2 x 2 contingency table, the (A, B, C, and D).There were no instances of Haplogroup X association of A or D Uneages with the Chumash and 46 Journal of California and Great Basin Anthropology { Vol. 26. No. 1 (2006)

Table 2 B or C Uneages with Uto-Aztecan groups is statisticaUy significant and relatively strong (x^ = 32.03, p < 0.001; DISTRIBUTION OF CALIFORNIA INDIAN MITOCHONDRIAL DNA LINEAGES ACCORDING TO HAPLOGROUPS T = 0.478). It would appear, therefore, based on the evidence of haplogroup frequencies alone, that these two Haplogroup groups had distinctively different population histories. Linguistic Grnup A B C D This becomes even more evident in the analysis of individual sequences reported below. Cltumashan Family Northern (Obispeiio) 0 0 0 3 Haplogroup A Lineages Central Purislmeiio 1 0 0 1 Haplogroup A was the least common among CaUfonua Ineseiio 3 0 0 2 Indian samples in our database. A total of thirteen haplo­ Barbareho 1 0 1 2 Ventureno 5 0 1 0 types were present among 17 Haplogroup A lineages Island (Cruzeiio) 1 0 0 0 (Table 3). Figure 5 Ulustrates how the CaUfomia Indian Chumashan Totals (11) (D) (2) (8) lineages in Haplogroup A are related phylogeneticaUy Holian Macro-Unit to each other m a network diagram. Most of these were Achumawi/Atsugewi 0 1 0 0 of Chumashan origin. At the center of the network are Esselen 1 D 0 0 found those samples recognized as the principal fotmding Salinan 3 2 0 1 Yuman-Cochimi Family haplotype for Haplogroup A in the Americas (AOl in Ipai 0 2 5 0 Fig. 5), caUed subhaplogroup "A2" by Forster et al. (1996). Tipai 0 0 1 0 Yuma 0 1 0 0 Several of the Chumash haplotypes are distinctive in Cochimf 0 0 1 G that they are differentiated from the presumed fotmding Hokan Totals (4) (B) (7) (1) haplotype by a T-> C transition at np 16093; a subset

Penutian Macro-Unit of these acquired a mutation that reversed the C-* T Wintuan Family transition at np 16111 that characterize virtuaUy aU other Wintu 0 2 0 G "A2" haplotypes in the Americas and some in northeast Southern Patwin 0 1 0 0 Asia (Schurr 2004:14; Tanaka et al. 2004:1841). Those Utian Family Sierra Miwok 0 3 0 1 Chumash A haplotypes with the T-^ C transition at np Southern Costanoan 0 1 2 1 16093 form a branching chain (see Figure 5). According Yokutsan Family Nim Yokuts 1 8 0 4 to the expectations of our model of in situ development Buena Vista Yokuts 0 0 1 2 combined with matrilocal residence, this pattem unpUes a Penutian Totals (1) (15) (3) (8) stable presence of Chumashan peoples within the region Uto-Aztecan Family over many miUennia, whereby mutations resulting in new Tubatulabal 0 2 1 1 haplotypes became fixed within the population. Numic Branch Non-Chumash lineages among our Haplogroup Kawaiisu 0 3 1 1 Mono 0 3 0 1 A samples were distributed as follows: Salinan (3), Takic Branch Esselen (1), Yokuts (1), and Luiseiio (1). The Esselen Gabrielino 0 1 0 1 Kitanemuk 0 0 2 1 and one of the SaUnan lineages represent the fotmding Serrano/Vanyume 0 2 1 1 haplotype in Figure 5. This SaUnan lineage originates Cahuilla 0 2 3 1 from the coastal region called Lamaca in the Mission Cupeno 0 1 1 G Luiseiio 1 4 13 G San Antonio records, which was adjacent to the Esselen. Uto-Aztecan Totals (1) (18) (21)

Ail California Indian Samples 17 44 4U 25 Yokuts sample and the single Luiseno sample represent ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnson / Lorenz 47

Salinan & Esselen Chumashan Yok-Utian Uto-Aztecan Yuman-Cochimi (N=7) (N=21) (N=24) (N=47) (N=10) A DB Figure 4. mtDNA Haplogroup distribution in Central and Southem California.

Table 3

CALIFORNIA INDIAN HVSl SEQUENCES FOR HAPLOGROUP A

CO OS CO 1612 9 1618 3 16183. 1 1618 9 1620 9 1622 1 1622 3 1625 7 1626 3 1627 8 Sample Etiinollnguistic Group Documenteil Origin Haplotype 1611 1 1629 0 1630 1 1631 1 1631 9 1632 7 1636 2

CRS T C G t - 1 T C C C T C C G T G C T

JJG39 Esselen Excelen AOl T • • . T • • • T • • A C JJ133 Salinan Isley, Lamaca district AOl T • • A c JJ135 Chumash Snajalayegua (Slinaxalyfwf) A02 X T • C A c JJ136 Chumash Nomgio {'Onomyo) A02 T • C A c JJ010 Chumash Sotonocmu {Soxtonokmu') AG3 C T • A c JJ07G Chumash Calahuasa {Kalawashaq') A03 C T • A c JJG91 Chumash Cheumen {Hichmn), S. Rosa 1. A03 C T • A c JJGG2 Chumash Mupu A04 C • ( A c 1 JJ153 Chumash Matilija {Mat'ilha) A05 C C ( A c JJ036 Salinan Sicpats A06 T • A c JJ040 Chumash Ventureiio (village not identified) A07 C T • A c JJG58 Tachi AGS T • (; C A X JJGB5 Chumash Miquigui {Miklw) AGS C T A A c JJ168 Chumash Cayegues {Kaymsh) A1G T • [ \ A c JJ221 Luiseno Pimixga {Pllmlchnga) All T • A r c JJ253 Chumash Sisolop (Shisholop) A12 c A c JJ123 Salinan (or Yokuts?) Mission S. Miguel (or S. Antonio?) A13 T • C • T • • . T • • A c 48 Journal of California and Great Basin Anthropology | Vol. 26. No. I (2008)

A02 Chumash Chumash A04&>

Salinan (or Yokuts?)

Chumashan Family Uto-Aztecan Family Hokan Stock Penutian Stock Chumash

Figure 5. Haplogroup A network diagram for California Indian mtDNA lineages based on HVSl sequences. the only Haplogroup A lineages from Yokutsan and do with the large number of Uto-Aztecan samples Uto-Aztecan groups in our database. Both Yokutsan and in our database (Table 2), which conforms to the Uto-Aztecan peoples are beheved to have spread into general predominance of haplogroups B and C among their ethnographicaUy documented territories sometime populations in the greater Southwest (Carlyle et al. 2000; between about four and a half millennia to about one Malhi et al. 2002, 2003; Merriwether 2002; Lorenz and and a half millennia before present (Golla, in press; Smith 1994,1996). Haplogroup B also had the greatest Moratto 1984). Our migration model predicts that during sequence diversity of any haplogroup in our database. this process, some lineages from the previously existing A total of 24 haplotypes occur among the 42 sequences populations would be incorporated into the incoming obtained for Haplogroup B (Table 4).^^ The founding groups, which were otherwise hkely to be characterized haplotype proposed for Haplogroup B among North by different haplotypes. We interpret the isolated A American Indian populations (Forster et al. 1996) is haplotypes found among Yokuts and Luiseiio populations represented by two samples, each from groups that spoke to be instances of this acquisition of older Uneages during languages within the proposed Penutian superfamily: the process of population replacement. Patwin and Sierra Miwok (Haplotype BOS in Figure 6). This fotmding haplotype is more prevalent elsewhere in Haplogroup B Lineages the Americas (MaUii et al. 2002). OiU" two Wintu samples The greatest number of lineages among our CaUfomia each represent a haplotype one mutational step removed Indian samples was characterized by Haplogroup B, from the founding Uneage (Haplotypes BOS and B15 in slightly more than Haplogroup C. This in part has to Figure 6). One of these exhibits an HVSl sequence that ARTICLE I Genetics, Unguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Uneages | Johnson / Lorenz 49

zsesi EK91

Qzegi 61E9L UCSL 86291 8iZ9L ttZZ9l 19291 ZfiZ9l EZZ9t 8LZ91

ZlZ9t C_D (U CJ C-D C_3 C_3 CJ) CD C_3 C_D CJ CU CJ C_3 C3 CLJ3 CJJ C_3 C_D CJ) C_D C_D CU CJ CJ) CJ CJ C_D CJD C_3 C_3 CJ3 C_3 C_3 C_3 CJ) C_D C_3 CJ> C_3 C_3 CU

EIZ91 C_3

68191 C_3 CJ3 C_3 CD CJ CJ C_3 C_3 C_D CU C_3 CJ) C_D C_D CJJ CO CJ) C_D CJ CJ CJ) C_3 C_3 CJ CJ) C_D C_D C_3 CD CD CD CD CD CO CD CD CJ3 CO CO CO CO CO mm •cc

mm •COCOCDCDCDCOCOCOCDCDCOCOCOCD CD* •CD-CDCDCO'CO- • -COCO* £8191 C_D CO CO CO CD CD CD CO CO CO CD CD CD CO CO CO CO CD CD CD CD CO CO CO CO CO CO CD CD CO CD CO CO CO CO CO CO CO CD CO CD CD

Z8191 CO CO CO CJ CO • CD trZ19l M UJ U ZgL9l a> 9 am

6Z191 iLigi 80191 E6091

ZGogi IZ091

•.— 1— T— CsJ CNI C-sl CNJ CNJ CNJ CNJ cn CO""^'<*-'«^lX^Lr3CDCDCOaDCO CO r— r— OO OD CD •>— C-^J CO -^a- LO CO I CO CD ^ 1— C-sJ CO *=»- CDCOCOCOCOCOCOCOCOCOCOCOCOCOCOCDCDCZIC^JCDCOCOCOCOCOC^CO''— •<— 1— 1— *— '— T— •<— 1— •,— CNJC-MC-sJtTsICNI CO CO CO CO oa CO CO GO CO CO CO CO CO CO CO ca cO CO CO CO CO CO CO CO CO CO CO CO CO GO CO CO CO CO CO CO CO CO CO CXD CX3 i-n

•= ca cfc^

cc ^ 53 : ti "^ S S^ a? . ' Q3 CTJ ro rri .

0! S CI S •pn O 0-- ^—• O 03 CJ >^ >-. 2-^ ^ ^ er t -^ =t cz Q3 ro C3 CD CO =3 1== C) CD CO OO a_ C_D Q_ ^ CO OQ -i: S GO

;?= .C3 i^ •£= JO :^

'ra 'ra "ro -^ eg 03 CO ro Q3 ^ S S 5 -^ :^ •-^ w t ca. S "= -CD ^ ° "ro ro CO CO CO c; -3 03 CD ^ CO , I CO OO CO Q_ . • -a: OO >- OO ; ^ S B a o -a _ ix:i ^aH -i^ ZSD :=2 .

1— ^ CNJ CD .,^ CO CD If? '— "^ ^ CD 1— ^ CO OO 1— LTD OJ CTsj OD ••— CD CO ^- CO •>— <^i LO cO i—-i CNI OD ,— CO t— CO COD T— CO I CNJ 1— r— oDLor^cxDi— CDcvic-sjcoi^-cDcvjcocococoLrD'.^i— oocNj-^d-i— coLocaDcocDrZoocoi— ^--OOCJDCJDCOOOCOCNI 50 Journal of California and Great Basin Anthropology | Vol 26. No. 1 (2006)

B21

Tubatulabal Cahuilla Luiseno, Wintu 319 Cahuilla g^gl

Tubatulabal

Mono 16362 811^^ Yokuts 16261 Patwin, B2oi 16298 ^^'Wok 16147 B16 16218 16319 16274 16184 ^^ B06 Salman Yokuts & B07 16093 1g311 802 & BIO Unidentified 16108 Unidentified 16343 Serrano)' California B01 809 16213C Luiseno, 1 16157 Indian ^^3^2 Cupenol6:|25 Yokuts Kawaiisu gQg 16223 B14l 16092 Wintu & Yuma Achumawi Bi9e 16129 Ipai B24( Mono Miwok r16»3 8 B22^P 16439 Uto-Aztecan Family

Mono Penutian Stock Hokan Stock

823 f I Costanoan ? Unidentified

Figure 6. Haplogroup B network diagram for California Indian mtDNA lineages based on HVSl sequences. is shared with the single Achumawi/Atsugewi lineage Westem Mono), and so the resultant star-like pattem is represented in our sample, illustrating some degree consistent with the hypothesized Yokutsan expansion of intermarriage among Northern California groups within the San Joaquin VaUey region and adjacent Sierra belonging to different linguistic phyla. Nevada foothiUs. The shared presence of the colonizing Twelve Haplogroup B samples distributed in six haplotype (B06 in Figure 6) among Yokuts, Sierra Miwok, haplotypes share a unique mutation that resulted in and Salinan Uneages probably derives from intermarriage a transposition of two nucleotides within the "poly- between the Yokuts and their neighbors. C" region. Instead of an A at np 16183 and a C at The non-Penutian portions of our Haplogroup np 16184, these two nucleotides reversed position so B Uneages are all associated with Uto-Aztecan and they contain an A->- C transversion at np 16183 and a Yuman groups. It is probably significant that there is no C^A transversion at np 16184 (Table 4). Haplotype overlap at all between the haplotypes represented by B06 contains only this transposition to distinguish it CaUfomia Penutians and those present among California from the fotmding haplotype B03. Five other haplotypes Uto-Aztecan (and Yuman) groups, implying separate branch from B06, each containing one or two subsequent population histories in geographicaUy separated regions. mutations (Figure 6). This star-like pattem is commonly It is difficult to discem phylogenetic pattems associated associated elsewhere in the world with migration events with particular Uto-Aztecan or Yuman subgroups in prehistory. All of the sequences containing the np because of the reticulations previously mentioned and 16183/16184 transposition are from Yokuts Uneages or because of past intermarriage between these groups, their immediate neighbors (SaUnan, Sierra Miwok, and who all practiced patrilocal residence. Nonetheless, ARTICLE I Genetics, Unguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnson / Lorenz 51 there are certain preliminary observations that can be CaUfornia Indian Uneages of uncertain origin. One of made. Although the frequently mutating np 16111 makes the interesting pattems revealed by the network diagram discernment of phylogenetic relationships problematic, it is the widespread phylogenetic relationships revealed is perhaps not coincidental that the three B haplotypes between neighboring Luiseiio and Ipai populations, even found among the Yuman lineages in our database all though they belonged to separate language famiUes. AU appear on the same branch among those containing five Ipai samples in Haplogroup C occur on three separate this mutation (Figure 6).^^ Also, there are suggestive "branches" that include either a shared haplotype with indications of at least one unique lineage among the their Luiseiio neighbors or are one mutational step UnguisticaUy and geographicaUy isolated Tubatulabal, one differentiated from them.This pattem is predictable based which is characterized by four mutations that distinguish on the documented intermarriage between these groups it from the founding B haplotype. Although the np 16111 and the long prehistory of Uto-Aztecan and Yuman mutational "hot spot" somewhat obscures its phylogenetic populations that were adjacent to one another in the position in Figure 6, this Tubatulabal Uneage is most Ukely Greater Southwest. Linguistic evidence of phonological related to a Desert CahuiUa Uneage with which it shares convergence between Takic and Yuman groups is entirely two markers (a T-> C transition at np 16092 and a C-*T consistent with the genetic data (Hinton 1991). transition at np 16147).^^ This provides a modicum of Two of the westem Takic groups, the Kitanemuk and support for the notion, based on Unguistic studies (GoUa the Vanyume (Desert Serrano), have unique haplotypes 2000b), that the Tubatulabal group was descended from a that warrant comment. One of the Kitanemuk sequences common Takic/Tubatulabal ancestral population (Haplotype C19) shares the T-» C transitions at np 16189 Four of the six Numic samples in Table 4, three and np 16311 that also have been identified among Kawansu and two Western Mono, are on a distinctive Northem Paiute populations in the Great Basin (Kaestle branch of their own, containing three haplotypes. Indeed, and Smith 2001; Malhi et al. 2003, 2004), suggesting a there is greater differentiation between these particular past interaction or shared history with Numic peoples, Numic lineages than an examination of their HVSl who Uved to the northeast of the Tehachapi Mountains sequence alone suggests. AU three diverge from other where the Kitanemuk homeland was located. The single Haplogroup B samples by not exhibiting the defining 9- Vanyume sample (Haplotyped CIO), with four mutations base pair deletion, even though they otherwise possess the not shared with any other Haplogroup C lineage, is quite distinguishing markers for Haplogroup B in their HVSl unusual. Two different individuals belonging to the same sequences. These samples were initially categorized matrUine participated in our study in order to double- as "other" in our restriction enzyme analysis, which check this distinctive pattern, and both sequences were was unexpected because of the unquestioned Native identical. The earUest female ancestor of this lineage was a American pedigree of those who provided the samples woman baptized at Mission San Femando, who had been to us. Subsequent examuiation of the HVSl sequences bom about 1750 in the vUlage of Topipabit, located along for these individuals revealed their Haplogroup B the Mojave River near the VictorviUe Narrows. Most afliUation. recently, mtDNA samples obtained from the teeth of two Late Period burials from a prehistoric cemetery near Haplogroup C Lineages PaUndale have yielded precisely the same HVSl sequence, Haplogroup C comes a close second to Haplogroup B demonstrating that this Uneage has a prehistoric presence in terms of its large proportion and diversity of Uneages in the Westem Mojave Desert region (Kemp, Eshleman, within our database. Table 5 Usts these 38 sequences, and and MaUii 2005).^* Its unique phylogenetic position within Figure 7 iUustrates the phylogenetic distribution of the Haplogroup C may suggest a greater antiquity for this 20 haplotypes identified within our California Indian haplotype (CIO) in the desert region, perhaps predating database. The long-recognized founding haplotype for the arrival of Uto-Aztecan populations. Haplogroup C is clearly apparent at the center of the Although Haplogroup C Uneages were overwheUn- diagram in Figure 7 (Haplotype C02); it occurred among mgly a Uto-Aztecan and Yuman-Cochimi phenomenon the Luiseiio and Kawaiisu, as weU as in two Southern among our samples, a few Haplogroup C haplotypes 52 Journal of California and Great Basin Anthropology | Vol. 26. No. 1 (2006)

^^ Cochimi, >v Luiseiio I^SQO 08 Luisefio & Cahuilla 15^ Cumash, 16051 , ^„ M6293 ^^ Ipai Yokuts, 07 16249 16825 0° Ipai Costanoan Ipai 11 03 16234 Luiseiio & Cahuilla 20 ( 16086 16235 Unidentified California Indian Coos 16189 16124 ,4 Unidentified 05^ California Indian Kitanemuk mvl & Tubatulabal Costanoan

Uto-Aztecan Family

Penutian Stock

Kitanemuk Hokan Stock Chumashan Family

Unidentified

Figure 7. Haplogroup C network diagram for California Indian mtDNA lineages based on HVSl sequences. were found among such disparate groups as the Yurok haplotype's dispersed distribution, and ancient DNA of Northwestern California, the Coos of the Oregon data from the regions in question wiU be necessary in Coast, the southem Costanoans, the HometwoU Yokuts, order to test its Ukelihood. and the coastal Chumash. One might suppose that the The haplotype represented by our single Yurok southem VaUey Yokuts and Chumash samples might be sample (C04 m Figure 7) is present in both a Barbareho derived from intermarriage with their Takic neighbors Chumash lineage and a Tipai Uneage fromnorthe m Baja to the south; however, it is more difficult to explain California, all from populations completely unrelated the presence of the same haplotype (C07) among the linguistically and widely separated from one another southem Costanoans. One possibUity would be that this along some 1000 kUometers of coastline. This haplotype particular haplotype derives from an "Old Uto-Aztecan" is defined by the lack of a T-> C transition at np 16325, presence in the San Joaquin VaUey prior to the Yokuts which furthermore has a broadly occurring distribution migration into the same territory, as has been proposed in the Northwest, Northeast, and Southwest regions of by Nichols (1988) and Moratto (1984:556). The presence North America (Malhi et al. 2002:910-911). Although of the C07 haplotype among the southern Costanoans, not currently recognized as a founding haplotype southem VaUey Yokuts, and Ventureiio Chumash could within Haplogroup C (Forster et al. 1996), it also is thereby resuU from intermarriage with the original Uto- present among Asian populations and is centrally Aztecan members of this lineage. This is but one among located in some network diagrams that cover the several conceivable hypotheses that could explain this North American continent (Malhi et al. 202:910).The ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnsor / Lorenz 53

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Table 6

CALIFORNIA INDIAN HVSl SEQUENCES FOR HAPLOGROUP D

csi^-ScM^-eooB^O'— EiSCco ^•*— ^c^>iesJesJc^>J^r^eomc*3CQm Sample EthnDlinguistic Group Documented Origin Haplotype CO ^'~> ^'~> ^'~' CO '•^•' fft ft CD CO CO CO CD

CRS G C G C A C C ( T G T r T

JJ006 Yokuts Hometwoli {Taneshach) 001 T • • • C c JJ025 Yokuts Tulamni 001 T c c JJ043 Yokuts Silelamne 001 T c c JJ047 Yokuts Yawdanchi 001 T c c JJ054 Yokuts Nutunutu 001 T c c JJ060 Tubatulabal South Fork, Kern River 001 T c c JJ098 Chunnash Tipu (Northern Chumash) 001 T c c JJ196 Chumash? Mission San Luis Obispo? 001 T c c JJ280 Mono Owen's Valley Paiute 001 T c c JJ363 Gabrielino Ouinquina (San Clemente Island) 001 T c c JJ059 Miwok So. Sierra Miwok 002 A T c c JJ163 Unidentified 002 A T c c JJ017 Kitanemuk Tejon region 003 T C c c JJ048 Kawaiisu Tehachapi region 003 T c c c JJ001 Chumash Miasap {Mi'asap) 004 T G 1r • ; c JJ067 Chumash Siujtu {Syuxtun) D04 T G 1r • ] c JJ076 Chumash? Calahuasa? {Kalawashaq') 004 T G 1r • ] c JJ165 Chumash Castait {Kashtayit) 004 T G 1r • ] c JJ050 Cahuilla Wavaaikiktum, Desert Cahuilla DOS T /\ c c JJ075 Costanoan Ensen D06 r T c c JJ093 Chumash Sucu {Shuku) 007 T 1r T c c JJ145 Vanyume Topipabit 008 T c JJ155 Yokuts Mission San Juan Bautista 009 T T c c JJ416 Salinan Lima 010 T • • T c c reported hypervariabihty of np 16325, however, prevents adjacent to Yokutsan peoples, leading us to propose geneticists from rejecting the competing hypothesis of that intermarriage accounts for the incorporation of convergence through independent mutations (Malhi et Haplogroup D Uneages into neighboring Uto-Aztecan al. 2002:915). populations, which are otherwise dominated by haplogroups B and C. However, the Island Gabrielino, Haplogroup D Lineages Desert Cahuilla, Barbareiio Chumash, and Salinan Haplogroup D is represented by 24 sequences distributed lineages (belonging to haplotypes DOl, DOS, D07, in 10 haplotypes within our database (Table 6). Ten and DIO, respectively), are likely to be independently of these samples (41.7 percent) belong to the widely derived and not accounted for by intermarriage with recognized founding Haplogroup D lineage occurring the Yokuts. The Island GabrieUno and Desert CahuiUa among American Indians (Forster et al. 1996). This lineages, in particular, and perhaps the Vanyume lineage haplotype (DOl) was present among the Yokuts (five as well (Haplotype DOS in Figure 8), may represent Uneages), Northem Chumash (two lineages),Tubatulabal, older matrilines incorporated among Uto-Aztecan Mono (Owen's Valley Paiute), and Island Gabrielino. groups during their expansion into California, as in With one notable exception, which we will discuss in the explanations previously proposed for the presence further detail below, most of the remaining haplotypes of Haplotype All among the Luiseiio (Figure 5) and are arranged about the basal Haplogroup D Hneage Haplotype CIO among the Vanyimie (Figiu-e 7). in a star-like pattern just one or two mutational steps One highly differentiated Uneage within Haplogroup removed from the founding type (Figure 8). Virtually all D is readily apparent in Figure 8. Considerable light of those haplotypes that are a single mutation removed has recently been shed on this haplotype, which occurs from the founding D haplotype are found in groups Uving prominently among western coastal and Santa Ynez ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Jotioson / Lorenz 55

Chumash Chumashan Family Yokuts Uto-Aztecan Family D09 Costanoan Penutian Stock D06 Hokan Stock

Unidentified

Yokuts, Mono, Tubatulabal, Gabrielino, Chumash

D03 Kitanemuk & Kawaiisu Miwok & Unidentified Cahuilla California Indian

Figure 8. Haplogroup D network diagram for California Indian mtDNA lineages based on HVSl sequences.

Valley Chumashan groups (our Haplotype D04). Four DISCUSSION samples exhibit the exact same pattern, which differs Mitochondrial DNA Uneages are useful for phylogenetic from the previously recognized founding haplotype studies because they descend only from a single parent, by possessing transitions A->- G at np 16241, C^T at one's mother, and therefore can be compared to each np 16301, and T^ C at np 16342, as weU as lacking the other to determine the closeness of genetic ancestry. The T-»- C transition at np 16325 (Table 6). A closely related simple branching diagram or "cladogram" that can be haplotype had previously been discovered by Rickards used to graph mtDNA phylogenies is not necessarily a and her coUeagues (1999) among the Cayapa Indians of good model to depict the real world of how language coastal Ecuador; it was proposed by these researchers that change occurs or how different peoples come together it represented an additional foundmg haplotype among to form descendant groups (Moore 1994). Nonetheless, American Indians. A sequence identical to that occurring when insights derived from linguistic prehistory and among the Chumash was recently discovered by Kemp archaeology are used in conjunction with genetic from an mtDNA sample obtained from a human tooth analyses, a new synthesis becomes possible regarding from an Early Holocene burial excavated by James Dixon past processes that resulted in changes in population in On Your Knees Cave (Dixon 1999:117-119; Kemp composition and ethnogenesis. Earlier in this paper, et al. in press). This finding supports the identification of we proposed that distinctively different genetic and this haplotype as being another founding Uneage for the linguistic patterns would result from different types of Americas. Other closely related Uneages have now been population interactions associated with migration events identified among Mexican populations, the Mapuche in prehistory. Our interpretations of the results of the Indians of southern Chile, and prehistoric peoples of mitochondrial DNA HVSl sequences are based on these Tierra del Fuego, among others (Kemp et al. in press). earUer predictions. We must recognize, however, that our 56 Journal of California and Great Basin Anthropology | Vol. 26. No. 1 (2006) sample sizes for California's various native populations coastal colonization model best matches the genetic are still rather modest, and that some undetermined diversity present among American Indian populations. amount of genetic diversity was undoubtedly lost during He proposes that transportation by watercraft would the nineteenth-century population decline. Therefore, encourage intermarriage outside of local groups and our observations are preliminary and wiU be subject to result in relatively rapid expansion beyond already settled modification as more extensive sampling and analysis areas (see also Erlandson 2002). Linguistic evidence for takes place in the future. the uniqueness of the Chumashan language family (GoUa, Our study has revealed some extremely interesting m press) and the genetic evidence presented here support results with regard to surviving Uneages that may descend the notion that coastal populations in the Santa Barbara from the uiitial migration events that settled North and Chaimel region could weU have been descended from this South America. The principal haplotypes present in those initial colonization event.^' fotmding populations have long been recognized (BaiUet Two significant clues exist in our mtDNA samples et al. 1994; Forster et al. 1996; Merriwether 2002; Schurr that Chumashan peoples had an ancient presence in 2004). Another founding haplotype has been detected south-central California. The first has to do with the within Haplogroup D, which is represented in CaUfomia aforementioned observation that the rare D haplotype by four of our Chumash lineages (Kemp et al. in press; (D04) is found primarily in Pacific coastal populations Rickards et al. 1999). This haplotype (D04) showed up on in discontinuous locations between British Columbia a distinctive branch four mutational steps removed from and the southernmost tip of South America (Kemp et the weU-known fotmding haplotype (DOl) in the network al. in press). The presence of this haplotype and some of diagram showing Uneage relationships within Haplogroup its derived Uneages among the Mapuche, Yahgan, and D (Figure 8). Now that this additional founding Imeage ancient peoples of Tierra del Fuego suggests that the has been discovered, what about other original haplotypes first wave of coastal migration might weU have included that might have lingered on among CaUfornia's native women representing this mtDNA founding lineage. Its peoples? A quick glance at Figure 7 shows that at least one presence among some interior groups in Mexico and other Uneage is isolated on a distant branch in its respective among prehistoric peoples in IlUnois could weU derive haplogroup diagram: CIO (Vanyume). This haplotype is from subsequent expansions of coastal peoples into the even farther removed ui the number of HVSl mutations interior of the North American continent. from the previously recognized founding haplotype for The second clue regarding the greater antiquity Haplogroup C than is the unconmion Chumash haplotype of Chumashan groups derives from the distribution of D04 within Haplogroup D One must consider, therefore, Haplogroup A, which is largely hmited to Northwest the possibility that this Vanyume lineage represents Coast and California groups inhabiting coastal regions an additional rare survival remaining from very early (Eshleman et al. 2004). Haplogroup A lineages are migrations that led to the peopUng of California. present ui their greatest abundance and diversity among Dixon (1999) has recently summarized the accumu­ Chumashan peoples, and the branching chain of Chumash lating evidence that favors a Pacific coastal migration route haplotypes Ulustrated ui Figure 5 is evidence of a lengthy by Paleoindian populations into the American continents and stable population throughout much of prehistory. by the end of the Pleistocene. Some anthropologists have Although both the rare D haplotype (D04) and the proposed that the great diversity of languages and language Haplogroup A Uneages could be derived from the same famiUes present along the Pacific Coast of North America coastal migration, we must also consider the alternative derives from such irutial colonization (Gruhn 1988,1992; hypothesis that they resulted from two independent Rogers 1985). On the basis of a sophisticated Imguistic populations becoming estabUshed and coalescuig in the analysis, Nichols (1992, 2000, 2002) has suggested that highly productive Santa Barbara Chaimel region. Indeed, by whatever route the contuient was mitiaUy populated, Nichols favors this scenario, based on her interpretations a subsequent coastal migration accounts for certain rare of a statistical analysis of the distribution of rare Unguistic features m languages distributed along the Pacific Coast features among American Indian languages (Nichols of the Americas. Fix (2002) has demonstrated that a 2002:287-290). Furthermore, Klar (2002) reports ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages {Jolinson / Lorenz 57 evidence of a non-Chumashan substrate m the Cruzeiio certain Great Basin and Plateau groups, and Eshleman language. Analysis of ancient mtDNA samples wiU be and Smith (m press) further note that the Takic distribution necessary to determine whether Haplogroup A and is similar to that of more ancient central California Haplogroup D Uneages have equal antiquity ui the Santa populations. The examination of specific haplotypes among Barbara region. these populations, however, shows that there is much Haplogroup A lineages characterized our single greater genetic distance between them than was suggested Esselen sample and three of our Salinan samples, by overaU sinularities in theU haplogroup distributions. suggesting that these could weU have persisted from the The migration of Uto-Aztecan groups into southem same early colonization along the Pacific shoreline that CaUfornia appears to have resulted in the introduction resulted m the estabUshment of a population along the of new genetic lineages mto the area, as weU as language Santa Barbara Channel (Eshleman et al. 2004; Eshleman replacement. At least one non-Yuman and non- and Smith, in press). The Unguistic evidence for ancient Chumashan language once existed in this region, as is contact between Esselen and the Chumashan languages demonstrated by a lexical and phonological substratum of (Shaul 1988), and the presence of Haplogroup A Uneages undetermined affUiation in Gabrielino (Bright and Bright from prehistoric burials in the Big Sur region (Eshleman 1976). Here and there, especiaUy in the desert areas and 2002:122), could signify that a Salinan language was not on San Clemente Island, there exist hints of the surviving mitiaUy spoken in the coastal region. It is possible that the mtDNA lineages of the earUer inhabitants of southem three Haplogroup A Uneages among oiu- SaUnan samples California. The distinctive Haplogroup C haplotype may have previously existed in the region prior to being discerned among the Vanyume; the rare presence of absorbed during a Salinan expansion towards the coast Haplogroup D lineages among the Vanyume, Desert under pressure from an incoming migration of Yokutsan Cahuilla, and Island Gabrielino; and the sole Haplogroup groups into the Central Valley region from the Great A sample from the most interior of the Luisefio viUages, Basui (see GoUa, in press). are probable survivals from the pre-Takic period. The Similarly, rare uistances of Haplogroup A lineages fact that these Uneages aU persisted in relatively marginal among the Yokuts and Luiseiio samples were likely areas suggests that the dryer, desert regions served as derived from earUer populations that were incorporated refugia for peoples who otherwise came to speak the into expanding groups who arrived later in these language of a dominant incoming group who co-opted respective regions (see also Eshleman and Smith, in more favorable habitats. press). The Yokuts Haplogroup A haplotype (A08) is Eshleman and Smith (in press) and GoUa (in press) particularly interesting because it is identical to a sequence have given thorough consideration to the evidence reported from a prehistoric burial in Esselen territory, pertaining to the timing and spread of language famiUes thus further suggesting an original coastal connection included within the Penutian macro-unit (see also for this haplotype (Eshleman 2002:122). Haplogroup Moratto 1984). Our data for Wintuan (3 samples) are stiU A lineages also have been identified among prehistoric too sparse to come to meaningful conclusions regarding populations living in marginal areas, such as the San the spread of this language family in Northem California, Clemente Islanders and the Pericii of the southem tip other than to note that at least one Wintu Uneage is shared of Baja California (Endicott et al. 2004; Potter 2004). with the linguistically unrelated Achumawi/Atsugewi. This pattern is fully consistent with the expectations Our sample is likewise too smaU to test the hypothesis of an initial coastal migration hypothesis, in which this that the Miwok-Costanoan (Utian) fanuly may have had haplogroup became estabUshed early among peoples with a longer presence in CaUfomia than the Yokutsan family; a maritune subsistence base and then persisted m certain however, we did find a genetic pattem that supports the coastal locations despite later population expansions.^" reconstruction of a relatively late expansion of Yokuts Based on an earUer subset of the data reported more peoples in the San Joaquin region with concomitant fuUy here (Tables 2-6), Eshleman et al. (2004) noted a absorption of older mtDNA lineages and intermarriage perceived similarity in haplogroup distributions between with neighbormg groups (see also Eshleman and Smith, the Takic and Yimian groups of southem CaUfomia and in press). 58 Journal of California and Great Basin Anthropology | Vol 26, No. 1 (2006)

Mitochondrial DNA research is coming into its °The available records pertaining to the sample classed as own as another means of exploring the prehistory of "Yuma" here are conflicting with regard to tribal affiliation. At CaUfornia's indigenous societies. Although our database least one record was seen that implied that the woman from southem Arizona may have been Papago instead. undoubtedly under represents the genetic diversity once present among the native peoples of Central and The Chukchansi sample might be of Choinimni origin instead. Southern California, the high-resolution genealogical The Island Gabrielino sample comes from the same matriline data do give us confidence regarding the pattems that to which Felicitas Montano belonged; she was one of John have been revealed to date. A coUaborative approach Harrington's GabrieUno consultants (McCawley 1996:17). utilizing the analytical techniques of human genetics and After GabrieUno, the Tataviam Icinguage had the next greatest taking advantage of California's rich ethnohistorical, number of speakers at Mission San Femando. People from these ethnographic, and Unguistic record has yielded and wiU two groups comprised 40 percent and 25 percent respectively of those baptized at the mission (Johnson 1997:252; Johnson and continue to yield insights into the prehistoric past of the Earle 1990). diverse peoples who uihabited the Pacific coast region. ^"When more than one sample was obtained from the same matriline, only one was included in our analysis to avoid dupUcation. NOTES '^^The HVSl sequences reported in this article have been Prediction 2 has been investigated elsewhere (Eshleman et al. submitted to the National Center for Biotechnology Information 2004) with regard to a wider geographical region extending for inclusion in the GenBank database (accession numbers beyond California. DQ383516-DQ383636). They are cross-referenced to the sample numbers appearing in Tables 3-6. These non-California Indian samples were often from family members or friends of donors who accompanied California ^^ The haplogroup affiliations of more than 100 of the 126 Indian participants in the study. Some samples descended from samples were determined tluough RFLP analysis. The remaining the Spanish-Mexican population of the Colonial Period were samples were assigned to haplogroups based on whether they derived from mtDNA lineages initially beUeved to be CaUfomia possessed the diagnostic HVSl markers for those particular Indian, but later determined to have originated ultimately haplogroups. HVSl sequences were obtained for 122 samples from a female ancestor who had immigrated to the region and out of the total of 126. whose female descendants had married a man who possessed California Indian ancestry. For example, one sample initially ^3 Smith et al. (1995) reported that a HVSl sequence from a believed to be Gabrielino was subsequently determined to single Porno individual belonged to Haplogroup X; however, have descended from a woman bom in Loreto, Baja California, genealogical documentation was not obtained pertaining to whose female lineage had likely originated in Sinaloa (Northrop this person's direct matemal ancestry. Further research will be 1987). That woman's great granddaughter married a Gabrielino necessary to determine if Haplogroup X Uneages exist among Indian man at Mission San Gabriel. So although the family was the Porno and whether they might yet be detected among other accurately identified as GabrieUno in the 1928-1933 California California native groups Indian enrollment records, the surviving mitochondrial DNA lineage was of Mexican Indian origin. ^^See Note 5 above.

^Further research will be necessary to locate contemporary ^^The number of haplotypes may be slightly less than Table 6 descendants of the single remaining Chumash mtDNA lineage indicates The difficulty in determining the number of Cs in the known to exist that has not yet been sampled for our study. This "poly-C" region between np 16180 and np 16193 in the mtDNA matriline descends from a woman bom on Santa Cruz Island sequence introduces some uncertainty in the assignment of and baptized at Mission San Buenaventura. haplotypes. We have reported the length variants in this HVl C-stretch in our tables according to the recommendations of "^See Milliken and Johnson (2005) for a discussion of village Wilson et al. (2002a, 2002b), but have eliminated positions locations. np 16182 through 16183.2 from consideration in creating the network diagrams in our figures, because the sequence variabiUty ^Sicpats may be the same rancheria (village) called "Gmimu" reported at these locations appears not to be phylogeneticaUy in Mission San Luis Obispo's registers. Both words apparently meaningful in all cases. meant 'carrizo place' in the Salinan and Northern Chumash languages respectively (Milliken and Johnson 2005:123). Thus, ^^The C->T transition at 16111 causes reticulation in the the Sicpats sample potentially could represent a Northern network diagram in Figure 6. This nucleotide position appears to Chumash lineage, even though it may be traced back to a be something of a "hot spot" in the mitochondrial genome, with woman baptized at Mission San Miguel, which was founded in evidence of having imdergone mutation within Haplogroup A southem SaUnan territory. also, among our California samples ARTICLE I Genetics, Linguistics, and Prehistoric Migrations: An Analysis of California Indian Mitochondrial DNA Lineages | Johnson / Lorenz 59

^^The T->C transition at np 16092 also appears in an 6pata REFERENCES sample and two Hispanic Uneages, also typed to Haplogroup B, in our larger database of Mexican American populations (to BaUlet, Graciela, Francisco Rothhammer, Francisco R. be reported elsewhere). It has also been reported for a sample Camese, Claudio M. Bravi, and Nestor O. Bianchi coUected in Chihuahua (Green, Derr, and Knight 2000). So the 1994 Founder Mitochondrial Haplotypes in Amerindian origin of this mutation may predate the entry of Uto-Aztecan Populations. American Journal of Human Genetics peoples into CaUfomia. The C-^ T transition at 16147 however, 54:27-33. appears only among our samples from CaUfomia. Bakker, Peter 2000 Rapid Language Change: CreoUzation, Intertwining, ^^See Eshleman and Smith (in press) for additional examples Convergence. In Time Depth in Historical Linguistics, of shared Uneages between modem individuals and preliistoric Colin Renfrew, April McMahon, and Larry Trask, burials elsewhere in CaUfomia. eds., pp. 585-620. Cambridge: McDonald Institute for Archaeological Research. ^^ Lacking CaUfomia evidence, Oppenheimer (2003:304) has opined that Paleoindian populations belonging to mitochondrial Bean, Lowell J. Haplogroup B may have expanded southward in an early 1978 Social Organization. In Handbook of North American migration along the Pacific coast. The data reported in our study, Indians, Vol. 8: California, Robert F. Heizer, ed., however, suggest that the earUest peoples more likely would pp. 673 -682. Washington, DC: Smithsonian Institution. have belonged to mitochondrial DNA haplogroups A and D BeUwood, Peter, and Colin Renfrew (editors) ^°See Dixon (1999:34-43) for a model of the different stages 2002 Examining the Farming/Language Dispersal Hypo­ of colonization of the Americas, spreading out from an initial thesis. Cambridge: McDonald Institute for Archaeological coastal zone of occupation. Research. Bettinger, Robert L, and Martm A. Baumhoff 1982 The Numic Spread: Great Basin Cultures in Competi­ tion. ylmen'canAnrig'Mify 47:485-503.

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