Mitochondrial DNA variation and the origins of the Aleuts

The mitochondrial DNA (mtDNA) variation in 179 Aleuts from five different islands (Atka, Unalaska, Umnak, St. Paul, and St. George) and Anchorage was analyzed to better understand the origins of Aleuts and their role in the peopling of the Americas. Mitochondrial DNA samples were characterized using polymerase chain reaction amplification, restriction fragment length polymorphism analysis, and direct sequencing of the first hypervariable segment (HVS-I) of the control region. This study showed that Aleut mtDNAs belonged to two of the four haplogroups (A and D) common among Native Americans. Haplogroup D occurred at a very high frequency in Aleuts, and this, along with their unique HVS-I sequences, distinguished them from Eskimos, Athapaskan Indians, and other northern Amerindian populations. While sharing several control region sequences (CIR11, CHU14, CIR60, and CIR61) with other circumarctic populations, Aleuts lacked haplogroup A mtDNAs having the 16265G mutation that are specific to Eskimo populations. R-matrix and median network analyses indicated that Aleuts were closest genetically to Chukotkan (Chukchi and Siberian Eskimos) rather than to Native American or Kamchatkan populations (Koryaks and Itel'men). Dating of the Beringian branch of haplogroup A (16192T) suggested that populations ancestral to the Aleuts, Eskimos, and Athapaskan Indians emerged approximately 13,120 years ago, while Aleut-specific A and D sublineages were dated at 6539 +/- 3511 and 6035 +/- 2885 years, respectively. Our findings support the archaeologically based hypothesis that ancestral Aleuts crossed the Bering Land Bridge or Beringian platform and entered the Aleutian Islands from the east, rather than island hopping from Kamchatka into the western Aleutians. Furthermore, the Aleut migration most likely represents a separate event from those responsible for peopling the remainder of the Americas, meaning that the New World was colonized through multiple migrations.     

Anthropometric variation among Bering Sea natives

Recent research indicates that anthropometrics can be used to study microevolutionary forces acting on humans. We examine the use of morphological traits in reconstructing the population history of Aleuts and Eskimos of the Bering Sea. From 1979 to 1981, W. S. Laughlin measured a sample of St. Lawrence Island Eskimos and Pribilof Island Aleuts. These samples included adult participants from St. George and St. Paul in the Pribilof Islands and from Gambell and Savoonga on St. Lawrence Island. The Relethford-Blangero method was used to examine the phylogenetic relationship between Aleuts and Eskimos. Anthropometric measurements for Native North Americans (measured by Boas and a team of trained anthropometrists in 1890-1904) and Native Mesoamericans (compiled from the literature for 1898-1952) were used for comparison. A principal components analysis of means for measurements and a neighbor-joining tree were constructed using Euclidean distances. All these tests revealed the same strong relationship among the focus populations. The R matrix from the Relethford-Blangero method clusters Aleuts and Eskimos separately and accounts for 97.3% of the variation in the data. Phenotypic variation within the population is minimal and therefore minimum F(ST) values are low. Genetic distances were compared to a Euclidean distance matrix of anthropometric measurements using a Mantel test and gave a high but not significant correlation. Our results provide evidence of a close phylogenetic relationship between Aleut and Eskimo populations in the Bering Sea. However, it is apparent that history has affected the relationship among the populations. Despite previous findings of higher European admixture in Gambell (based on blood group markers) than in Savoonga, Savoonga has greater within-group variation in anthropometric measurements. Anthropometrics reveal a close relationship between Gambell and St. Paul as a result of European admixture. The St. George population was the most divergent of the populations, indicating that it diverged from the Eskimos and St. Paul because of the compounding effects of genetic drift and limited European gene flow. These findings are in agreement with previous anthropometric and genetic studies of the Aleut and Eskimo populations and support the utility of anthropometrics in inferring population history and structure.     

Genetic variation in the Aleuts of the Pribilof Islands and the Eskimos of Kodiak Island

A sample of Aleuts residing in the Pribilof Islands of St. Paul (N = 163) and St. George (N = 62) and Eskimo residents of Kodiak Island (N = 294) have been typed for genetic variation at 31 discrete genetic markers. Of these, 16 were polymorphic and 15 were monomorphic. Several private polymorphisms previously reported in Eskimo or Alaskan Amerindian populations were absent in both the Aleuts and Kodiak Island Eskimos. Genetic distance analysis shows considerable genetic differentiation between Aleuts and Kodiak Island Eskimos.     

HLA genes in populations of the Aleutian islands

We typed a subset of the Aleut population for HLA loci (HLA-A, HLA-B, HLA-DRB1, HLA-DQB1) to obtain an HLA profile, which was compared to other Eurasian and Amerindian populations for studying Aleut origin and its significance on the peopling of the Americas. Allele frequencies at the four loci were identified in an Aleut sample using standard indirect DNA sequencing methods. Genetic distances with Amerindians and Eurasians were obtained by comparing Aleut allele frequencies with a worldwide population database (13,164 chromosomes). The most frequently extended HLA haplotypes were also calculated. We also generated Aleut relatedness dendrograms and calculated correspondence relatedness in a multidimensional scale. Both neighbor-joining dendrograms and correspondence analysis separated Aleuts from Eskimos and Amerindians. Aleuts are closer genetically to Europeans, including Scandinavians and English. Our results are concordant with those obtained by Y-chromosome analysis, suggesting that most male Aleut ancestors of our sample came mainly from Europe.     

Analysis of mitochondrial DNA diversity in the aleuts of the commander islands and its implications for the genetic history of beringia

The Aleuts are aboriginal inhabitants of the Aleutian archipelago, including Bering and Copper (Medny) Islands of the Commanders, and seem to be the survivors of the inhabitants of the southern belt of the Bering Land Bridge that connected Chukotka/Kamchatka and Alaska during the end of the Ice Age. Thirty mtDNA samples collected in the Commanders, as well as seven mtDNA samples from Sireniki Eskimos in Chukotka who belong to the Beringian-specific subhaplogroup D2, were studied through complete sequencing. This analysis has provided evidence that all 37 of these mtDNAs are closely related, since they share the founding haplotype for subhaplogroup D2. We also demonstrated that, unlike the Eskimos and Na-Dene, the Aleuts of the Commanders were founded by a single lineage of haplogroup D2, which had acquired the novel transversion mutation 8910A. The phylogeny of haplogroup D complete sequences showed that (1) the D2 root sequence type originated among the latest inhabitants of Beringia and (2) the Aleut 8910A sublineage of D2 is a part of larger radiation of rooted D2, which gave rise to D2a (Na-Dene), D2b (Aleut), and D2c (Eskimo) sublineages. The geographic specificity and remarkable intrinsic diversity of D2 lineages support the refugial hypothesis, which assumes that the founding population of Eskimo-Aleut originated in Beringan/southwestern Alaskan refugia during the early postglacial period, rather than having reached the shores of Alaska as the result of recent wave of migration from interior Siberia.     

Skeletal evidence of health and disease in pre-contact Alaskan Eskimos and Aleuts

There have been relatively few paleopathological studies of arctic populations to date, compared to other regions of North America. Studies aimed at elucidating patterns of health and disease in arctic peoples prior to contact and assessing inter- and intraregional differences in disease patterns have been particularly few. In the present study, five pre-contact skeletal samples (N = 193), representing 4 Eskimo populations from northern coastal Alaska and 1 Aleut population from the eastern Aleutian Islands, were examined macroscopically for the following indicators of health status: cribra orbitalia, porotic hyperostosis, trauma, infection, dental caries, abscesses, antemortem tooth loss, periodontal disease, and dental attrition. In addition, archeological and epidemiological data were used to help reconstruct the health of these populations. The goals of the analysis were 2-fold: 1) to assess the pre-contact health of North Alaskan Eskimos and Aleuts in order to provide a baseline comparison for the post-contact health of these groups, and 2) to determine if any differences in disease patterns exist between the Eskimos and Aleuts that might be related to differences in their physical environment, subsistence patterns, and cultural practices. The analysis revealed that both groups suffered from a variety of health problems prior to contact, including iron deficiency anemia, trauma, infection, and various forms of dental pathology. Statistical comparisons of the 2 groups revealed that Eskimos and Aleuts had different patterns of health and disease prior to contact. Most notably, the Aleuts had a significantly higher frequency of cranial trauma and infracranial infection than the Eskimos, while the latter had a significantly higher frequency of enamel hypoplasia. An examination of the physical and cultural environment of the 2 groups reveals several possible explanations for these differences, including warfare, subsistence pursuits, and housing practices. The documentation of these differences indicates that variability in pre-contact disease patterns can be identified between hunter-gatherer populations living in similar environments and exhibiting similar general lifestyles. Am J Phys Anthropol 107:51–70, 1998. © 1998 Wiley-Liss, Inc.     

Origins of linguistic diversity in the Aleutian Islands

The Aleut language, currently spoken along the Aleutian chain and the Pribilof and Commander islands, is the only language in its branch of the Eskimo-Aleut language family, and traditional methods of linguistic reconstruction have neither satisfactorily explained its relationship with languages on the Asian continent nor its development from Proto-Eskimo-Aleut. Linguistic reconstruction has always been important in understanding the prehistory and history of the Aleuts, and new approaches in comparative linguistics, more comprehensive information on typological features of neighboring languages, and continuing language documentation allow us to propose a rich and continuous history of contact with various groups of people. I evaluate evidence that the Aleut language may have been shaped by contact with neighbors in Asia and Alaska, eventually giving rise to its differentiation from the Eskimo languages. I look at dialect differentiation along the Aleutian chain and what this differentiation reveals about the migration trends of the Aleut along the chain. I look at the colonial expansion of the Aleut-speaking area and resulting additional varieties of Aleut in the historical period. Finally, I review the effects of the Russian and American colonial periods on the Aleut language and the severe endangerment that the language faces today as a result. I conclude that there is evidence of possible Aleut contact with both neighboring peoples; however, much of this evidence has not yet been subjected to systematic comparative reconstructions. Linguistic evidence supports theories of at least two westward expansions of Aleuts along the island chain, but it is not yet clear what motivated the dialect differentiations. Finally, I offer some thoughts on directions for future dialect studies and the continuing documentation of Aleut.     

Physical growth of St. Lawrence Island Eskimos: Body size,proportion, and composition

Growth patterns of body size, proportion, and composition were analyzed in 57 male and 56 female Eskimos from St. Lawrence Island in the Bering Sea, ranging in age from 1.23 through 19.82 years. Age-groups means for whites and blacks of the U.S. Health Examination Survey served as reference data. Relative to HES data, the Eskimo sample were shorter with lower values for leg length, while there were no differences from the reference values for sitting height. The Eskimos also had higher values of Quetelet's Index, the sitting height/height ratio, and the upper arm muscle circumference, while there were no differences in body weight or triceps skinfold thickness. Differences from the reference data were more pronounced in males than in females. The growth patterns for size and body proportion are in conformity with known relationships between morphology and climate.     

Dental chipping in Aleuts, Eskimos and Indians

Analysis of dentitions belonging to 324 prehistoric and protohistoric Aleuts, Eskimos and northern Indians, all of whom were regularly meat-eaters, reveals a significant difference between Eskimos and Aleut-Indians for a little known type of tooth wear. This wear is characterized by severe crushing and/or flaking of the crown surface of one or more teeth (termed “pressure-chipping”). It occurs chiefly in dentitions of high arctic Eskimos of Alaska, Canada, and Greenland, and significantly less often in the teeth of Kodiak Island Eskimos, Aleuts and northern Indians. Sex differences do not exist but pressure-chipping occurs significantly more often in adult (21–x years) than in non-adult (0–20 years) Eskimos. The exact mechanism(s) responsible for pressure-chipping is unknown, although ethnographic accounts of Eskimo eating habits suggest that crushing of hard substances such as bone was involved. The severity of this wear could have contributed to the selection for, or preservation of, large and complex crowns in high arctic Eskimos. Pressure-chipping is offered as evidence favoring the view that tooth size (longevity) may have had in the past some adaptive value.     

Distribution of albumin variants Naskapi amd Mexico among Aleuts, Frobisher Bay Eskimos, and Micmac, Naskapi, Mohawk, Omaha, and Apache Indians

In order to help define the boundaries of the distribution of the albumin variants Naskapi and Mexico which are polymorphic among several American Indian groups, we examined sera from Micmac, Mohawk, Northwest River Naskapi, Omaha and Apache Indians, and from Aleuts and Eskimos. Sera from a total of 1,524 individuals were examined. Using a cellulose acetate membrane electrophoretic system with Tris-Citric acid at pH 5.4 we were able to distinguish normal albumin and both variants in the same run. Naskapi and Mexico variants were absent from Aleut, Eskimo, Micmac, Mohawk and Omaha samples. The albumin Naskapi variant was present in an allele frequency of 0.03 in the Naskapi Indian sample. Albumin variants Naskapi and Mexico were found in the Apache sample at frequencies of 0.016 and 0.037, respectively. This report supersedes that previously published by Schell and Agarwal ('76). Generally, within an area there is a correspondence between changes in the frequency of albumin variants and changes in the ethnic background and history of the area's populations. At the same time, when viewing widely separated areas, relationships between distant groups based on linguistic and cultural similarities are paralleled on a biologic level by the distribution of normal albumin and variant albumins.     

Origins of Aleuts and the genetic structure of populations of the archipelago: molecular and archaeological perspectives

We summarize the results of a field and laboratory research program (1999-2006) in the Aleutian Islands on the origins of the inhabitants of the archipelago and the genetic structure of these populations. The Aleuts show closest genetic affinity to the contemporary Siberian Eskimos and Chukchi of Chukotka and differ significantly from the populations of Kamchatka (the terminus of the archipelago) and Alaskan Eskimos. Our findings support the hypothesis that the ancestors of the Aleuts crossed Beringia and expanded westerly into the islands approximately 9,000 years ago. The Monmonier algorithm indicates genetic discontinuity between contemporary Kamchatkan populations and western Aleut populations, suggesting that island hopping from Kamchatka into the western Aleutian Islands was highly unlikely. The primary determinant of the distribution of genes throughout the archipelago is geography. The most intimate relationship exists between the genetics (based on mtDNA sequences and intermatch/mismatch distances) and geographic distances (measured in kilometers). However, the Y-chromosome haplogroup frequencies are not significantly correlated with the geography of the Aleutian Islands. The underlying patterns of precontact genetic structure based on Y-chromosome markers of the Aleut populations is obscured because of the gene flow from Russian male colonizers and Scandinavian and English fishermen. We consider alternative theories about the peopling of the Americas from Siberia. In addition, we attempt a synthesis between archaeological and genetic data for the Aleutian Islands.     

Morphologic characteristics of the Alaskan Eskimo dentition. II. Carabelli's cusp

A study of coastal and inland Alaskan Eskimos revealed that a faintly developed Carabelli's cusp was present in 42.7%. No sex difference in the occurrence of Carabelli's cusp was evident and no family showed any difference in the distribution of the frequencies when each family was compared to the rest of the population. A general tendency toward a higher prevalence of Carabelli's cusp in the coastal Eskimos may be due to more admixture with white people along the coast. Alaskan Eskimos have a significantly higher frequency of Carabelli's cusp than do Aleuts. Statistical evaluation revealed that in the Alaskan Eskimo dentition the formation of Carabelli's cusp is independent of the size of the molars and the suppression of the third molars.     

A burial cave in the western Aleutian Islands, Alaska

During the 1998 field season, the Western Aleutians Archaeological and Paleobiological Project (WAAPP) team located a cave in the Near Islands, Alaska. Near the entrance of the cave, the team identified work areas and sleeping/sitting areas surrounded by cultural debris and animal bones. Human burials were found in the cave interior. In 2000, with permission from The Aleut Corporation, archaeologists revisited the site. Current research suggests three distinct occupations or uses for this cave. Aleuts buried their dead in shallow graves at the rear of the cave circa 1,200 to 800 years ago. Aleuts used the front of the cave as a temporary hunting camp as early as 390 years ago. Finally, Japanese and American military debris and graffiti reveal that the cave was visited during and after World War II. Russian trappers may have also taken shelter there 150 to 200 years ago. This is the first report of Aleut cave burials west of the Delarof Islands in the central Aleutians.     

Skeletal growth of children from the Iron Age site at K2 (South Africa)

Cross-sectional growth data were obtained from the skeletal remains of children from the Iron Age site of K2 near the Limpopo River. Standard measurements of the diaphyseal lengths of the long bones from both limbs were recorded and compared to published skeletal data. For this purpose, data on Eskimo and Aleut skeletons, Libben skeletons, and skeletons from Indian Knoll and Altenerding were used. An attempt to study growth allometrically was made. K2 children were growing as well as children from these other groups. Comparison of data for K2 children with those on living South African “Cape Coloured” rural children, studied during the late 1980s, shows the similarity of growth of both groups. © 1996 Wiley-Liss, Inc.     

The St. Lawrence Island Eskimos: genetic variation and genetic distance

The Eskimos of St. Lawrence Island have been typed for genetic variation at 44 discrete genetic loci. Three private polymorphisms, at the 2,3-diphosphoglycerate mutase, peptidase B, and purine nucleoside phosphorylase loci, have been observed, which may be useful in future studies of genetic relationships between Eskimos and other circumpolar populations. Genetic distance analysis reveals a close relationship between the St. Lawrence Island Eskimos and other Eskimo populations and that the Eskimo populations form a distinct cluster from Amerindian populations. The St. Lawrence Island Eskimos appear to be more similar to Asiatic Eskimos than to other groups. Caucasian admixture in this population is estimated to be between 2 and 7%.     

Three-rooted mandibular first permanent molars and the question of American Indian origins

Three-rooted mandibular first molars (3RM1) are characteristic of Asian and Asian-derived populations, particularly Aleuts (whose 3RM1 frequency is the highest in the world) and Eskimos. Similarities in the frequency of these teeth between American Indians and contemporary peoples of southeastern Asia indicate a closer relation between these groups than between American Indians and Aleut-Eskimos. Three-rooted mandibular first molar frequency does not differ significantly in males and females except in Aleut-Eskimos. Bilateral asymmetry of 3RM1 is relatively frequent in both sexes and all groups. All American Indian groups examined have a low frequency of 3RM1 pointing to a single Asian origin, except Athabaskan-speaking Arizona Navajos, whose 3RM1 frequency approaches that of Aleut-Eskimos. There is no evidence at present of any significant local microevolution of 3RM1 in two testable prehistoric American Indian groups, although genetic drift had possibly occurred in a few series of 3RM1-deficient southwestern U. S. prehistoric Western Pueblo Indians. No adaptive value can be found for 3RM1 in Indians. In prehistoric western U. S. Indians geographic frequency variation is only slightly greater than the very slight (and non-significant) testable temporal variation. Three migrations from Asia seem best to explain New World 3RM1 frequency variation.     

New approaches to the pre- and post-contact history of Arctic peoples.

The last few years have witnessed the addition of new techniques and research strategies to the study of the population history of Arctic peoples. Osteon-photon analysis of bone cores provides an improved method of assigning age at death to skeletons. Consequently, it is possible to improve calculations of life expectancy and relate them to pathological correlates such as osteoporosis, separate neural arches, spina bifida and arthritis along with regular growth changes. This capability enables much better utilization of pre-contact skeletons and therefore of the numbers, density and composition of populations before European contact. The general picture emerging from skeletal studies, census records and living populations is, in Arctic Eskimos, one of high fertility, high mortality and short length of life, with a slow population growth rate. Aleuts show lower fertility, lower mortality and longer length of life, also with a low population growth rate.     

Genetic studies of low abundance human plasma proteins. II. Population genetics of coagulation factor XIIIB.

Plasma samples from a large number of different ethnic groups, consisting of U.S. whites, U.S. blacks, Eskimos from Kodiak and St. Lawrence Island, Aleuts of the Pribilof Islands, and three Amerindian groups from Canada and Mexico have been analyzed by isoelectric focusing followed by immunoblotting to determine the magnitude of genetic variation at the F XIIIB structural locus. The synthesis of published data and our new data demonstrate remarkable variation in the distribution of the three common alleles at this locus and establishes F XIIIB as an extremely informative marker for population differentiation and evolutionary studies. Genetic distance analysis based on this variation separates Caucasian, black, and Mongoloid populations into three distinct clusters.     

Analysis of contingency table data on torus mandibularis using a log linear model

The hypothesized sexual difference in the incidence of torus mandibularis in Eskimoid groups, as well as age and group differences, was studied in two Northwest Territories Eskimo groups and in an Aleut group (examined by Moorrees). The data were analyzed using statistical methods new to the field of anthropology. The method analyzes data sampled from a multinomial distribution. A hierarchy of log linear models are fitted to the cell counts of a contingency table. An iterative proportional fitting procedure is used to obtain expected cell counts under each log linear model. The “goodness of fit” of each model is tested by the log likelihood ratio. This statistic can be partitioned into additive components such that differences between models can be tested. In this way a “best” model, from the hierarchy of models, is chosen. Among these three groups of Eskimos, the incidence of torus mandibularis was not affected by sex but was affected by age and was different between the three groups.     

Morphologic characteristics of the Alaskan Eskimo dentition. III. Number of cusps on the upper permanent molars

A study of 35 coastal and 64 inland Alaskan Eskimos revealed a reduction in the number of cusps from the first to the third maxillary molar. While 97% of the first molars had four cusps, only 39.6% of the second molars and 15.2% of the third molars had that number. The reduction occurs through elimination of the hypocone. No statistically significant sex difference in the trend towards reduction in the cusp numbers was found. In the inland female group the occurrence of four cusps in the maxillary second molar was statistically higher than in the coastal female group. This may be due to a more pronounced racial admixture of white people along the coast. A similar difference, although not statistically significant, was found in the corresponding male groups. Alaskan Eskimos have a tendency towards a lower frequency of four cusps on all three maxillary molars than Aleuts. Only the second molar exhibited a statistically significant difference in this respect. A statistical evaluation revealed that in the Alaskan Eskimo maxillary first and third molars the reduction of cusps is independent of the size and form of the molars and of the suppression of the third molar. For the second molar, however, the groups with four well-developed cusps showed significantly larger buccolingual diameter.