,

Acta Anthropogenetica, Vol. 8 (1 & 2): 79-109 (1984)

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. I

'

Peopling of Northern North America: Clues from Genetic Studies

EMOKE J.E. SzATHMARY

Department of Anthropology, McMaster University, Hamilton, Ontario, Canada

Abstract The paper reviews the archaeological evidence for the length of human occupation in N. America and raises the question whether single or multiple movements of people out of Asia into America occurred, pointing out that considerable genetic variation can occur in small isolated populations in relatively short periods of time. The entire subarctic culture area is populated by speakers of either Athapascan or Algonkian language families. The archaeologic record for tracing the origin of these linguistic groups depends on items of material culture· and these have been used to trace the origin of the modern peoples back for a few thousand years. Comparison between groups based on genetic data suffers from unevenness of the data for various Athapascan-and Algonkian-speaking groups. The problem is made more difficult by the smallness of populations and inadequate sample size. The gene diversity measure H of Nei has been used on data for the A i:hapaskan Dogrib. It suggests that there was probably significant gene diversity present in sub-arctic groups in pre-contact times. Probably this is true also for the Algonkians as typified by the Ojibwa. Examination of the apportioned gene diversity shows that the bulk of the diversity exists within groups rather than between groups. Genetic clues to the peopling of the Americas derive from specific marker genes and from genetic distance statistics. The distribution of the Di 6 and the Gmza;boast alleles suggest that Athapaskan genetic links are towards the Bering Sea area while Algonkian ·connections are towards the south. Nei's genetic distance statistic was calculated for 13 populations using 14 blood group and enzyme loci. The dendrogram derived from the D matrix shows that Eskimos and Chukchi cluster together, and the Athapaskans are closer to the Eskimos than are the Algonkians. These relationships could be valid if the origin of Eskimos goes back to a population of Asiatic Beringia and that populations north of the late Wisconsin ice sheets included a group that led to the Athapaskans whilst populations south of the Wisconsin ice sheet led to the Algonkians.

Introduction To many, the recency of the occupation of the Americas and the relative sameness of its peoples in traits of superficial morphology suggest

•

Szathmary

that little can be learnt by studying Amerindian biology. Archaeologists, for example, who extol "the richness and variety of prehistoric American cultl.lres" and marvel at the linguistic diversity on the two continents 42 seem unaware that considerable biological heterogeneity also exists in the Aniericas. What appears, on the surface, to be "a relatively uniform racial composition'"4 2 masks a considerable amount of genetic variability63 64 92 • Furthermore, even the superficial "general sameness" 10 refers only to features such as pigmentation of the skin, hair and eyes, h3ir form, the scantiness of facial and body hair, and dental traits. On the other hand, characteristics such as stature or craniofacial form and d\n'lensions do show variation in both North and South America 10 is 45. What general interest most Americanists have in Amerindian biology is reflected: in questions concerning the physical type of the original immigrants 44. There is also some concern whether the diversity that is evident in their descendants resulted from successive population increments from Asia, or whether this was produced by evolutionary forces within the New World. Understandably, questions of the first sort are best satisfied by skeletal data-which, for the requisite time periods of first occupation, are vanishingly rare 113• However, genetic data obtained from the descendants can a1so be used to establish genetic relationships and then: are many examples known where "genetic distances reflect not only ethnology but also history and archaeology" 21 76. While it is a fact that the question of Amerindian origins and affinities has long fascinated Americanists and continues to do so (e.g., Laughlin and Barper 51 ), this is not the only appeal the peoples of the Americas have for human biologists. Focus on genetic relationship, which is but one cause of genetic similarity distracts attention from the ·obverse-namely, genetic difference. The vast amount of genetic diversity in humans discovered by the application of improved biochemical detection methods begs explanation. Nee\fft has been the most eloquent of all geneticists in articulating the importance of Amerindian studies-particularly on: those groups that have m1intained biological and sociocultural integrity-for making inferences about the evolutionary forces that main.tain genetic variability in populations. This conviction has had the practical result that some South American tribal groups are better known-genetically, linguistically, and sociologically than most nation states. Studies on North American Indians have not been a~ detailed, in part because of the disruption most groups have experienced from post Columbian immigrants. Nevertheless, it remains one of the tasks of human biologists to quantify the existing genetic differences within and between these populations as well. In addition, although the difficulties may be more severe, human

Peopling of Northern North America: Clues from Genetic Studies

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81

biologists must provide explanations for how these differences arose and Rre maintained. Such explanations can be sought by testing the observed data against various models supplied by population genetics theory. The approaches woulg yield answers specific for the people of North America. However, their ultimate purpose for the human biologist would be to achieve a deeper understanding of how the evolutionary process operates on the human species. This paper provides a general framework for the interpretation of Amerindian biologic variability. It also supplies some examples of recent studies on the estimation of genetic diversity within groups of subarctic Amerindians. Lastly, it shows how such research can lead into deduction about Amerindian affinities with each other and with Asians.

Amerindian Biology : Current Perspectives Few people growing up in North America have escaped the adage "if you've seen one Indian you've seen them all." The statement from which this maxim evolved has been attributed to Antonio de Ulloa, a Spanish scientific traveller writing in the year 1772. In 1951 Stewart and Newman93 documented how this opinion, espoused as it was by the most eminent men of "early" anthropology, served to retard our understanding of Amerindian biological diversity. They concluded that (I) notions of Amerindian homogeneity, (2) the attribution of Amerindian variability to waves of migration, or (3) reluctance to interpret Amerindian diversity with the methods and theory of population genetics represented a failure of Americanists to keep apace with developments in the biological sciences. In their opinion "data, old and new, will require interpretations set within / the broad framework of modern biology, with special emphasis on population genetics, ~ystematics and prehistory" (Stewart and Newman, 93 p. 34). This dictum was certainly observed by Brues10 in her treatment of Amerindian variability. She accepted as the baseline in her initial formulation a single archaeologically derived date (15,000 years BP) for first human entry into North America. With some simple calculations Brues showed that even with a very slow rate of population increase it would have been possible for a group of 100 "founders" to have 100 million descendants in 10,000 years. She saw no need to postulate mass migrations, certainly not to explain the distribution of culture traits, and argued that with the exception of the Eskimos and Aleuts the "genetic make-up of the populations of the Americas was largely determined by the first migrants." Diversification of this slowly expanding group, probably through the actions of genetic drift and natural selection produced what variability exists today.

82

Szathmary

The accuracy of Brues' interpretation is less dependent on the evolutionary mechanisms she postulated than on the archaeological conteitt into which the interpretation must be cast. These include at minimum a correct estimate of the first occupation of the Americas, as well as the availability of a detailed, continents-wide prehistoric record. Nothing of this sort currently exists, hence the archaeologically derived baselines which form the initial assumptions for the biological model, are subject to modification and possibly even reversal. This can be illustrated by contrasting the framework Brues employed with some recent developments in archaeology. Brues assumed an initial time depth of 15,000 years BP-a conservative, non-controversial estimate available more than a decade ago. This time depth meant entry into the New World towards the end of the climax of the Late Wisconsin glaciation, when the Bering land bridge was at its greatest extent. By 13,000 to 14,000 BP the Bering Strait bad reappeared, and by 11,800 BP it became an unfordable barrier between Asia and America "to any except skilled boatmen" 43 • Accordingly, the isolation of the first Americans from Old World contacts occurred within one or two thousand years after arrival of the "founders". Their southern expansion coincided with the retreat of ~he glaciers. The new prehistoric evidence shows that humans were present as far east as the Old Crow basin of Canada's Yukon Territory as early as 27,000 years ago 44• Furthermore, it can no longer be doubted that people were present south of the Late Wisconsin ice sheets by 21,070 BP (Meadowcroft Rock Shelter, 40°17"N, 80°29"W 1 • How these hunting bands got south of the ice is a biologically important issue, as well as a matter of some archaeological controversy. The pivotal problem is whether it was always possible for groups to move from Beringia to the central Great Plains, or whether the coalescence of the Cordilleran and Laurentide ice sheets effectively isolated the people south of the ice from those north of the ice for several thousand years. If the former possibility existed, then Brues' model needs little revision beyond a greater time depth. If the latter is correct, then there may indeed have been different "waves of migration" into and across the Americas-not necessarily the Paleoindian equivalent of organized convoys, but certainly, as the ice retreated, the gradual southward (and northward) movement of peoples that differed to some degree genetically. The evidence for an ice-free corridor-out ofBeringia along the Mackenzie River and continuing south along the eastern foothills of the Rocky Mountains is accumulating. Detailed geomorphologic, climatic, and floral and faunal studies provide environmental reconstructions that suggest

Peopling of Northern North America : Clues from Genetic Studies

83

strongly that the corridor was inhabitable for periods of time between 25,000-15,000 BP 30 • However, the so-called "ice-free" corridor was not open continuously. Fladmark30 suggests that ice between latitudes 55°-60°N probably blocked the corridor at the height of the glacial climax in west central North America, that is, between 18,000-i5,000 BP. In such a case people north of the ice were separated from those to the south by a minimum of 3000 years. While such a period may be insignificant geologically 44 it is of some consequence for genetic differentiation. Three thousand years approximates 120 generations. Not an insignificant amount of genetic difference can accumulate in small populations isolated from each other for such lengths of time.

Indians of Subarctic North America: (a) LINGUISTIC RELATIONSHIPS The current distribution of indigenous people in Canada and Alaska shows that the entire subarctic culture area (Figure 1) is populated by speakers of either Athapaskan or Algonkian Amerindian languages. Athapaskan languages are confin::d to the west, while Algonkian languages are spoken in the central and eastern regions107 • From where and when did the Athapaskans aud Algonkians come to inhabit such a vast territory? Algonkian and Athapaskan are distinct language families, with no known connections between them 12 • Neither language family is confined to the subarctic, but by far the greatest number of Athapaskan tongues are spoken in the north, while the greatest number of Algonkian languages are spoken south of the subarctic107 • Opinions concerning the origin of these people are mixed. Linguistic analysis suggests the ancient homeland of Athapaskan speakers was probably located in Alaska4s, in its eastern interior and may have ex.tended a considerable distance into Canada49 . Modern descendants of these ancestral Athapaskans speak a minimum of 11 languages in Alaska and likely another 12 in Canada. Greater precision regarding the exact number of languages is not possible, because Athapaskan, like many other language families, is a language and a dialect complex in which it is not always simple to determine boundaries 48 49 . Current classification of the Algonkian languages lists 26108 , although earlier works included only 17 6 • The problem here also is the difference between a dialect complex and a distinct language. Algonkian languages, like Athapaskan, are not confined to the subarctic culture area. However, while all extra-subarctic Athapaskans are thought to be descendants of people who had moved south from the northwestern region of the continent49, the same cannot be said for Algonkian. The central and eastern

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Peopling:or Norther~North America: Clues from Genetic Studies

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subarctic currently includes the dialect conti11uum Cree-Montagnais with numerous dialects between 25 , as well as Ojibwa in north-western Ontario. The latter, however, is part of the Ojibwa-Potawatomi continuum that extends south of the western Great Lakes. All other Algonkian languages are spoken, and other dialect continua are located south of the subarctic culture area reaching as far west as the American Rocky Mountains and along the eastern seaboard as far south as the Carolinas. Identification of the ancient homeland of Algonkian-speakers has not been done with the linguistic criteria employed by Krauss for Athapaskans -that is, the region of greatest linguistic diversity. However, Siebert89 has argued cogently that reconstruction of proto-Algonkian words from floral and faunal word lists of several Algonkian languages, and the correlation of these with ancient ranges and climates places the proto-Algonkian homeland in a restricted region of south central Ontario, between Lakes Huron and Ontario. This suggests that Algonkians are of a southern derivation, unlike the Athapaskans. (b)

ARCHAEOLOGICAL RELATIONSHIPS

The archeological record is not particularly helpful in addressing questions of ultimate origin raised by current language distributions. The 'in-situ' hypothesis prevails, whereby living peoples are thought to be the lineal descendants of people whose prehistoric remains are located in the regions the living now occupy. Because skeletal data are either completely missing from the ancient time periods (e.g., Alaska), or are rare (e.g., Archaic samples from the Great Lakes :75 ), and because such data are almost non-existent for historic subarctic people&9 s 103 the archaeologic record depends completely on items of material culture. These have been used successfully to trace the modern peoples back in time for a matter of a few thousand years. Thus, the earliest sequence thought to show continuity in lithics with any historic Algonkian is the Laurel tradition (earliest date 200 BC) of northwestern Ontario 24 110 111 • For Athapaskans, agreement among archaeologists stops for sequences earlier than A.D. 100025 • The roots of the earliest Algonkian and Athapaskan traditions are thought to lie in the various Archaic cultures that antecede them in the subarctic area25 36 • Some of these in turn are believed to have developed out of the earlier Paleo-Indian big-game-hunting stage identified by the use of various kinds of lanceolate points. Wrightl 10 and Harp 36 for example, both agree that Paleo-Indians from the central Plains (Plano tradition) moved north and southeast as the Laurentide glaciers retreated, ultimately to give rise to the Shield Archaic people (5000 BC-1000 BC).

86

Szathmary

Wright has suggested, on the basis of the continuity he sees between lithics and inferred 1ifeways between Shield Archaic and ensuing Laurel culture, that the Shield Archaic people may have spoken an Algonkian language 11 e>. Harp, however, while seeing "considerable merit" in this argument, thinks that the Shield Archaic was the foundation for all Indian groups which had a "basic taiga economy" and :this includes Athapaskan as well as Algonkuan peoples30 • It is worth noting that Dumond 25 in his summary of the Northern Archaic tradition ( 4000-2000 BC), more commonly thought to be ancestral to Athapaskans does not derive them from a Paleo-Indian base. Rather, he notes that others have tried to link the Northern Archaic sequence with the earlier occurring indigenous Alaskan Paleo-Arctic tradition, even though the continuity between the two traditions is not satisfactorily demonstrated. Accordingly, Dumond repeats the conventional view that the Northern Archaic reflects a forest-adapted tradition that was carried northward· by migrants "after the end of the Pleistocene glaciation" 25 • The Northern Archaic is linked, by virtue of the most commonly occurring artifact in it-the side-notched point-to the Archaic of eastern North Americal? 5 • This Archaic stage, called the Boreal (earliest sites c. 4000 BC) by Harp 36 and Laurentian, by Wright 111 , does not appear to have any connection to an earlier Paleo-Indian projectile point tradition. Nevertheless, Harp thinks the Boreal Archaic "gave rise to the early cultures of the Quebec-Labrador peninsula" -Montagnais and Naskapi, both groups bein ~ Algonkian-speakers. Wright110 111 , who also sees the Boreal (=Laurentian) Archaic as very different from the Shield Archaic, suggests that the former represents a second Archaic population with a different technology and "different antecedents". What existed between Boreal and Shield Archaic peoples may have been interaction based on trade, much as existed between historic Algonkians and Iroquoian-speakers of the northeast. Implicitly in Wright's view, these Laurentian ( =Boreal) Archaic people were not ancestors of the Algonkians. I think it is safe to say that given the sparseness of the evidence, it is risky indeed to draw correspondences between the direct antecedents of historic peoples and the earlier Archaic cultures. Thus, "although it seems eminently logical to assume that it* evolved into the various Indian cultures that occupied the region at the time of European discovery" 36 indubitable continuities are not established between Archaic and subsequent traditions for any part of the subarctic culture area. Furthermore, the validity of inferring ethnicity from stone tools is *i.e., Archaic

Peopling of NJrth~rn North America : Clues from Genetic

Studi~s

87

questionable. Tools have ::runctions. Pursuit of similar subsistence activities may require stone tools of similar size and shape (e.g .• reflecting a "taiga economy" 36 ) ,in which case similarity does not indicate anything about linguistic or biological relationship. Setting these difficulties aside, were the various archaeological speculations accepted, then all subarctic Indians would be derived from two interior Indian groups. The Algonkians would stem ultimately from Paleo-Indians who moved northeast from the region of the central Great Plains, with a possible increment in the east from people of the pre-projectile point stage. The latter moved north with the spread of the forests. The same group is thought to have led to the appearance of the Athapaskans across the continent in Alaska and adjacent Canada. (c) A HYPOTHESIS OF POPULATJON RELATIONSHIP A more critical observer might state, as has Dumon 0

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Table IV.

Distribution of the Im:nunoglobulin Gene Gmza;b03st in some Siberian and native American populations

Language Family and Population

Location

Chukotan Reindeer Chukchi Inland Chukotka Coast Chukchi Ctukot Peninsula Eskimoan Eskimo New Ch:tplino, Siberia St Lawrence Island, U.S. North Alaska Igloolik, Canada Thule, W. Greenland East Greenland Athapaskan Arctic Village & Ft. Kutchin Yukon, Alaska Dogrib NWT, Canada Alberta, Canada Chipcwayn Navajo Southwestern U.S. Algonkian Pikangikum, Ontario N. Ojib\\a Wikwemikong, Ontario S.E. Ojibwa Alberta Cree Uto-Aztecan New Mexico, U.S. Zuni Papago New Mexico, U.S. Arirona, U.S. Pima South American Central and South America 28 tribes

Gmza.bOlst

Reference No.

.153 .099

94 94

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94 29 59 60

.143

59

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104 83 83

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106 106 94

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85

less, there is no evidence at all in any of the gen~tic traits known for these people that sugg~sts they differ from more western subarctic Algonkians 1M. Montagnais and Nasbpi differcnc~s are with the Athapaskam 98 • In sum, the ecologicll data suggest that selection, were it operating on Gmza;boM and Dia, should have op::rated in the same direction in both the Athapaskan and Algonkian occupied subarctic regions for approximately ~imilar lengths of time. Nevertheless the distribution patterns of the&~ genes difftr between these pe\>ples.

Peopling of Northern North America : Clues from Genetic Studies

99

To invoke chance as the mechanism whereby the different distribution p1tterns of Gmza;bo 3 st and Dia arose in the subarctic zone is to provide no explanation. The prob1bility th1t drift of Dia produced a pattern shared by 10 Indian groups on the one hand from which 6 Indian groups on the other hand differ, and that this difference should correspond to known linguistic differences is very remote. The same could be said about the discreteness of the Gmza;bo3st distribution. What is more plausible is that the basic differences in the distribution of thes~ genes reflect different population origins. Jf so, then clearly Athapaskan genetic links are towards the Bering Sea area, while Algonkian connections are towards the south. (ii) THE EVIDENCE FROM STUDIES OF GENETIC DISTANCE Assessment of genetic similarities between populatiom are commonly done with statistics that measure genetic distance. This is less a contradiction than it seems, for the smaller the distance the greater the genetic similarity. The advantage that distance statistics have over information provided by the distribution of markrr genes, is that the d:stance measures can make use of all genetic data available in common for the populations compared. Thus, genes whose frequencies differ in quantitative fashion, and whose frequencies on scrutiny do not necessarily fall into any easily detectable patterns are nevertheless informative when used in conjunction with many such genes.

•

The statistic that has been used most often to detect genetic differences among northern North American and Asian population is Nei'& 67 standard distance, D. Szathmary 99 has also used Cavalli-Sforza and Edwards' 13 chord distance measure, and found a significant and very strong positive correlation between the two statistics (see also Jorde 47 ). This was an interesting finding, for neither the mathematics nor the theoretical formulations of the evolutionary process in the two approaches are the same. Cavalli-Sforza and Edwards13 assume that evolutionary divergence between populations over time is modulated chiefly by the processes of genetic drift and selective drift-that is, natural selection operating differently in different places and times. Nei's67 63 aµproach, however, assumes that populations diverge through time through the accumulation of new and different mutations in each of the descendant groups. Accordingly, Nei's standard distance D, for protein and enzyme loci, measures the number of codon differences per locus that are detectable by the laboratory methods used to determine genotype-electrophoresis . When blood group genes are used in the computation of D, the statistic loses its biological meaning. However, it still provides a statistical

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ALAStr races of man, Caucasoids, Negroids and Mongoloids. Am J Hum Genet 1974, 26: 421-443. 70 Nei M and Roychoudhury AK. Genetic relationship and evolution of human races. Evolutionary Biology, Vol. 14, ed. Hecht MK, Wallace Band Prance GT, pp. 1-60, Plenum Press: New York, 1982. 71 Nei M, Tajima F and Tateno Y. Accuracy of estimated phylogenetic trees from molecular data. II. Gene frequency data. J Mol Evol 1983, 19 : 153-170.

Peopling of Northern North America: Clues from Genetic Studies 72 73

74 75 76

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78

79

80

81

82 83

84

85

86 87 88

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Emo~e J.E. Szathmary Department of Anthropology McMaster University Hamilton, Ontario Canada