Australian aborigines represent the first branch from Eurasian antecedents: odontometric evidence

Most genetic data suggest that Australian aborigines and Southeast Asians associate, but their relative evolutionary relationship has remained obscure. Historically, the study of tooth crown variables has been important in establishing phylogenetic relationships. Through the quantification of whole tooth structure (GDP), including root, pulp, and enamel, a likely Eurasian phylogeny emerged from a canonical discriminant analysis of the microevolution among the populations. The analysis suggested that in modern human evolutionary history, Australian aborigines are the best representative extant population (first branch) from an unknown antecedent Eurasian founder population. The next branch from the Asian-based antecedent population was Caucasoids. Within the resident antecedent East Asian population, Southeast Asians then evolved, followed by a branch that lead to antecedent east Central Asians. Mongolians and all Native Americans independently evolved from this antecedent east Central Asian population. The relatively short morphogenetic separation between two areas that have been isolated for great periods of time, i.e., Australian aborigines and Native Americans, suggests that their association is not due to gene flow.     

Metric dental variation of major human populations

Mesiodistal and buccolingual crown diameters of all teeth recorded in 72 major human population groups and seven geographic groups were analyzed. The results obtained are fivefold. First, the largest teeth are found among Australians, followed by Melanesians, Micronesians, sub-Saharan Africans, and Native Americans. Philippine Negritos, Jomon/Ainu, and Western Eurasians have small teeth, while East/Southeast Asians and Polynesians are intermediate in overall tooth size. Second, in terms of odontometric shape factors, world extremes are Europeans, aboriginal New World populations, and to a lesser extent, Australians. Third, East/Southeast Asians share similar dental features with sub-Saharan Africans, and fall in the center of the phenetic space occupied by a wide array of samples. Fourth, the patterning of dental variation among major geographic populations is more or less consistent with those obtained from genetic and craniometric data. Fifth, once differences in population size between sub-Saharan Africa, Europe, South/West Asia, Australia, and Far East, and genetic drift are taken into consideration, the pattern of sub-Saharan African distinctiveness becomes more or less comparable to that based on genetic and craniometric data. As such, worldwide patterning of odontometric variation provides an additional avenue in the ongoing investigation of the origin(s) of anatomically modern humans.     

Dental and cranial affinities among populations of east Asia and the Pacific: the basic populations in east Asia, IV.

The origins of the four major geographical groups recognized as Australomelanesians, Micronesians, Polynesians, and East and Southeast Asians are still far from obvious. The earliest arrivals in Sahulland may have migrated from Sundaland about 40,000-50,000 years B.P. and begun the Australomelanesian lineage. The aboriginal populations in Southeast Asia may have originated in the tropical rain forest of Sundaland, and their direct descendants may be the modern Dayaks of Borneo and Negritos of Luzon. These populations, the so-called "Proto-Malays," are possible representatives of the lineage leading to not only modern Southeast Asians, but also the Neolithic Jomon populations in Japan. The present study suggests, moreover, that the Polynesians and western Micronesians have closer affinities with modern Southeast Asians than with Melanesians or Jomonese.     

Population prehistory of East Asia and the pacific as viewed from craniofacial morphology: The basic populations in East Asia,VII

Distance analyses were applied to 11 craniofacial measurements recorded in samples from East and Southeast Asia, Australia, Melanesia, Polynesia, and Micronesia for the purpose of assessing the biological affinities and possible origins of these populations. A clear separation between Australomelanesians and other populations from East and Southeast Asia and the Pacific is evident. The craniofacial variations suggest that the generalized Asian populations (Negritos, Dayaks, Lesser Sunda Islands, etc.) represent at least part of the morphological background of not only the majority of present Southeast Asians, but also the Neolithic Jomon people and their lineage in Japan, Polynesians, and western Micronesians. The original craniofacial features of Southeast Asians may have occurred as the result of convergent microevolution due to similar environmental conditions such as tropical rain forest. This supports the local-evolution hypothesis for modern Southeast Asian craniofacial features. © 1993 Wiley-Liss, Inc.     

Evolutionary relationships of human populations on a global scale

Using gene frequency data for 29 polymorphic loci (121 alleles), we conducted a phylogenetic analysis of 26 representative populations from around the world by using the neighbor-joining (NJ) method. We also conducted a separate analysis of 15 populations by using data for 33 polymorphic loci. These analyses have shown that the first major split of the phylogenetic tree separates Africans from non-Africans and that this split occurs with a 100% bootstrap probability. The second split separates Caucasian populations from all other non-African populations, and this split is also supported by bootstrap tests. The third major split occurs between Native American populations and the Greater Asians that include East Asians (mongoloids), Pacific Islanders, and Australopapuans (native Australians and Papua New Guineans), but Australopapuans are genetically quite different from the rest of the Greater Asians. The second and third levels of population splitting are quite different from those of the phylogenetic tree obtained by Cavalli- Sforza et al. (1988), where Caucasians, Northeast Asians, and Ameridians from the Northeurasian supercluster and the rest of non- Africans form the Southeast Asian supercluster. One of the major factors that caused the difference between the two trees is that Cavalli-Sforza et al. used unweighted pair-group method with arithmetic mean (UPGMA) in phylogenetic inference, whereas we used the NJ method in which evolutionary rate is allowed to vary among different populations. Bootstrap tests have shown that the UPGMA tree receives poor statistical support whereas the NJ tree is well supported. Implications that the phylogenetic tree obtained has on the current controversy over the out-of-Africa and the multiregional theories of human origins are discussed.      

Chimpanzees as an outgroup for the examination of human dental evolution

In the assessment of human origins, chimpanzees (Pan troglodytes, henceforth called Pan) represent the best hominoid outgroup for comparisons. Such an outgroup roots the "anatomically modern" human population cluster, or continuum. This study incorporates chimpanzees into a worldwide modern human database of quantified complete tooth variables (approximately 30 per tooth; e.g., root, pulp, enamel) in an attempt to develop a more accurate phylogeny of the hominoid continuum, with only intervening extinct hominids missing. Canonical discriminate analysis was performed mainly among Liberian common chimpanzees and global samples of humans. The first canonical variable explained 70% of the total variance and showed a tight cluster of humans, with chimpanzees as a distant outgroup. Within the human community, first non-San Bushman, sub-Saharan Africans and Andamanese, and then, close in, Australian aborigines were positioned towards Pan. Their relative orientation suggested an African human origin with the first branch within sub-Saharan Africa: sub-Saharan Africans and San Bushmen. Next, Andamanese Negritos, and then Australian aborigines, formed the early first surviving modern human lineage to leave Africa. Thin enamel and big teeth with relatively large roots characterized Pan nonmolar teeth. Humans showed a generalized sexual dimorphism for all teeth, with males having bigger teeth, bigger relative roots, and thinner enamel than females, while only Pan canines had significant and impressive sexual dimorphism. Interestingly, Pan molars were not larger than human molars. The data suggest that although hominids underwent two dental macroevolutionary events, the lineage leading to modern humans only experienced anterior tooth-size reduction. The suggested evolutionary significance of the observed total tooth variation is discussed.     

Heterochronic quantitative microevolution: Dental divergence in aboriginal Americans

Studies of tooth crown morphology alone have proven valuable in defining human population differentiation. We test the hypothesis that quantitative comparisons of more complex whole tooth structure may prove informative in understanding human diversity. Three disparate populations in Native American genetic history were compared: Kodiak Island Western Eskimos, Peruvian Inca Amerindians, and Southeast Asians. Enamel depth (an increasing gradient extended from Southeast Asians to the Inca) and root parameters were the most discriminating variables. The observed microevolution appears to result from variation in timing of known X-linked, Y-linked, and autosomal genes that affect either ameloblast or odontoblast differentiation. The dental traits were sexually dimorphic, the effect being more pronounced in aboriginal Americans, with male teeth having robust roots and thin enamel compared to female. Southeast Asians were isometrically related. The prominence of sexual dimorphism and the importance of sex-linked genes in the determination of the dental phenotypes suggest that sexual selection was one evolutionary force acting on early Asian populations. Subsequently, the selection appears to have been relaxed in Southeast Asians. Observed divergence of tooth shape among the populations, i.e., differences in the appropriation process of tooth primordia, was mainly the consequence of genetic drift modulating heterochronic regulators of homeotic genes. © 1996 Wiley-Liss, Inc.     

Late archaic and modern Homo sapiens from Europe, Africa, and Southwest Asia: Craniometric comparisons and phylogenetic implications

The craniometric affinities among Neandertals. Upper Palcolithic Europeans, early anatomically modern Southwest Asians, and archaic and modern Africans are investigated using univariate and multivariate methods. For the first time, it is possible to analyse the North African finds Dar-es-Soltane 5, Nazlet Khater, and Wadi Kubbaniya. It was not possible to include the Neandertals from Central Europe due to their poor state of preservation. The results point to, first, a basic distinction between Neandertals on the one hand and modern humans from all geographic regions on the other, and, secondly, to great similarities between modern African and European populations. Late archaic sapiens specimens from Africa were more similar to Upper Paleolithic Europeans than were the Neandertals. The results do not support the hypothesis that a regional evolution giving rise to modern humans took place in Europe. The results are, however, consistent with the hypothesis that modern populations originated in Africa and spread to Europe from there.     

Dental perspectives on the population history of Southeast Asia

This article uses metric and nonmetric dental data to test the "two-layer" or immigration hypothesis whereby Southeast Asia was initially occupied by an "Australo-Melanesian" population that later underwent substantial genetic admixture with East Asian immigrants associated with the spread of agriculture from the Neolithic period onwards. We examined teeth from 4,002 individuals comprising 42 prehistoric and historic samples from East Asia, Southeast Asia, Australia, and Melanesia. For the odontometric analysis, dental size proportions were compared using factor analysis and Q-mode correlation coefficients, and overall tooth size was also compared between population samples. Nonmetric population affinities were estimated by Smith's distances, using the frequencies of 16 tooth traits. The results of both the metric and nonmetric analyses demonstrate close affinities between recent Australo-Melanesian samples and samples representing early Southeast Asia, such as the Early to Middle Holocene series from Vietnam, Malaysia, and Flores. In contrast, the dental characteristics of most modern Southeast Asians exhibit a mixture of traits associated with East Asians and Australo-Melanesians, suggesting that these populations were genetically influenced by immigrants from East Asia. East Asian metric and/or nonmetric traits are also found in some prehistoric samples from Southeast Asia such as Ban Kao (Thailand), implying that immigration probably began in the early Neolithic. Much clearer influence of East Asian immigration was found in Early Metal Age Vietnamese and Sulawesi samples. Although the results of this study are consistent with the immigration hypothesis, analysis of additional Neolithic samples is needed to determine the exact timing of population dispersals into Southeast Asia.     

Dental anthropological indications of agriculture among the Jomon people of central Japan. X. Peopling of the Pacific

The high rate of crown caries (8.6%; 119/1,377 teeth) and other oral pathologies in 101 central Japan Middle to Late Jomon Period (ca. 1000 B.C.) crania indicate a level of carbohydrate consumption consistent with an agriculture hypothesis. Because Jomon dental crown and root morphology shows strong resemblances with past and present Southeast Asians, but not with ancient Chinese or modern Japanese, Jomon agriculture could be of great antiquity in the isolated Japanese islands. These dental data and other assembled facts suggest that ancestral Jomonese might have carried to Japan a cariogenic cultigen such as taro before the end of the Pleistocene from tropical Sundaland by way of the now-submerged east Asian continental shelf.     

Genetic diversity and evidence for population admixture in Batak Negritos from Palawan

Anthropologists have long been fascinated by the isolated hunter-gatherer populations in Southeast Asia (SEA) collectively known as "Negritos." However, the origins and affinities of these groups remain unresolved. Negritos are characterized by their short stature, dark skin color, and wiry hair, and they inhabit the Philippines, Malay Peninsula, and the Andaman Islands. Among Philippine Negritos, the Batak are of particular interest in understanding population interactions in the region due to their location on Palawan Island, which likely formed a corridor by which human migrations entered the rest of the Philippine archipelago from Island SEA. Here, we extend current understanding of the distribution of genetic diversity in Negritos by presenting the first analysis of mitochondrial DNA and Y-chromosome diversity among the Batak. We show that the Batak are genetically distinct from Negritos of the Andaman Islands and Malay Peninsula and instead bear most resemblance to geographically proximate Philippine Negritos and to non-Negrito populations from the Philippines and Island SEA. An extensive degree of recent admixture between the Batak and their neighbors is indicated by the high frequency of recently coalescing haplogroups in the Batak that are found throughout Island SEA. The comparison of results from these two loci further lends support to the hypothesis that male-biased admixture has, in particular, been a prominent feature of the interactions between the Batak and surrounding non-Negrito populations.     

Evolution of beta-globin haplotypes in human populations

The beta-globin haplotypes of 852 chromosomes from 12 populations in the Asia-Pacific region are described. These data are combined with those from other populations in an investigation of the affinities of regional human populations. Both partial maximum-likelihood and distance Wagner methods indicate that Africans are the most divergent group, with the remaining populations branching in the following order: Australian Aborigines, Highland Melanesians, Lowland Melanesians, Indonesians and Micronesians, Polynesians, east Asians, Indians, and Europeans. This pattern of relationship is consistent with that indicated by other data. Analysis of the evolution and distribution of haplotype occurrence provides some limited support for an origin of modern humans in Africa. Otherwise, however, it was not useful in further elucidating the evolutionary history of human populations.     

Denisova admixture and the first modern human dispersals into Southeast Asia and Oceania

It has recently been shown that ancestors of New Guineans and Bougainville Islanders have inherited a proportion of their ancestry from Denisovans, an archaic hominin group from Siberia. However, only a sparse sampling of populations from Southeast Asia and Oceania were analyzed. Here, we quantify Denisova admixture in 33 additional populations from Asia and Oceania. Aboriginal Australians, Near Oceanians, Polynesians, Fijians, east Indonesians, and Mamanwa (a "Negrito" group from the Philippines) have all inherited genetic material from Denisovans, but mainland East Asians, western Indonesians, Jehai (a Negrito group from Malaysia), and Onge (a Negrito group from the Andaman Islands) have not. These results indicate that Denisova gene flow occurred into the common ancestors of New Guineans, Australians, and Mamanwa but not into the ancestors of the Jehai and Onge and suggest that relatives of present-day East Asians were not in Southeast Asia when the Denisova gene flow occurred. Our finding that descendants of the earliest inhabitants of Southeast Asia do not all harbor Denisova admixture is inconsistent with a history in which the Denisova interbreeding occurred in mainland Asia and then spread over Southeast Asia, leading to all its earliest modern human inhabitants. Instead, the data can be most parsimoniously explained if the Denisova gene flow occurred in Southeast Asia itself. Thus, archaic Denisovans must have lived over an extraordinarily broad geographic and ecological range, from Siberia to tropical Asia.     

Mitochondrial DNA variation in Nicobarese Islanders.

The aboriginal populations living in the Nicobar Islands are hypothesized to be descendants of people who were part of early human dispersals into Southeast Asia. However, analyses of ethnographic histories, languages, morphometric data, and protein polymorphisms have not yet resolved which worldwide populations are most closely related to the Nicobarese. Thus, to explore the origins and affinities of the Nicobar Islanders, we analyzed mitochondrial DNA (mtDNA) hypervariable region 1 sequence data from 33 Nicobarese Islanders and compared their mtDNA haplotypes to those of neighboring East Asians, mainland and island Southeast Asians, Indians, Australian aborigines, Pacific Islanders, and Africans. Unique Nicobarese mtDNA haplotypes, including five Nicobarese mtDNA haplotypes linked to the COII/tRNA(Lys) 9-bp deletion, are most closely related to mtDNA haplotypes from mainland Southeast Asian Mon-Kmer-speaking populations (e.g., Cambodians). Thus, the dispersal of southern Chinese into mainland Southeast Asia may have included a westward expansion and colonization of the islands of the Andaman Sea.     

Dental morphology of the early Hoabinian, the Neolithic Da But and the Metal age Dong Son civilized peoples in Vietnam

The early Hoabinian, the Neolithic Da But and the Dong Son (early Metal age) civilized peoples in northern Vietnam were investigated based on dental morphology and were compared with specimens from surrounding Northeast and Southeast Asia including Australians and Melanesians. In both the metric and nonmetric tooth traits, the Hoabinian and Da But specimens had dental features similar to the prehistoric Southeast Asians and the Australo-Melanesians, but also had partially Northeast Asian characteristics. On the other hand, the Northeast Asian features become distinct in the dentition of the Dong Son people, which have close ties with the modern Vietnamese. Thus, the Vietnamese, as well as the other modern Southeast Asians and Japanese, are considered to be a blend of indigenous Southeast Asians who are closely related to the Australo-Melanesian lineage, and migrants from Northeast Asia.     

Out of Africa with regional interbreeding? Modern human origins

A central issue in paleoanthropology is whether modern humans emerged in a single geographic area and subsequently replaced the preexisting people in other areas. Although the study of human mitochondrial DNAs supported this single-origin and complete-replacement model, a recent paper(1) argues that humans expanded out of Africa more than once and regionally interbred. However, both the genetic antiquity and the impact of the African contribution to modern Homo sapiens are so great as to view Africa as a central place of human evolution. Despite the possibility that out-of-Africa H. sapiens interbred with other populations, this evidence is more consistent with the uniregional hypothesis than the multiregional hypothesis of modern human origins.     

Early Holocene human skeletal remains from Santana do Riacho,Brazil: implications for the settlement of the New World

In this study we compare the cranial morphology of several late Paleoindian skeletons uncovered at Santana do Riacho, Central Brazil, with worldwide human cranial variation. Mahalanobis Distance and Principal Component Analysis are used to explore the extra-continental morphological affinities of the Brazilian Paleoindian sample. Santana do Riacho is a late Paleoindian burial site where approximately 40 individuals were recovered in varying states of preservation. The site is located at Lagoa Santa/Serra do Cipó, State of Minas Gerais. The first human activities in this rockshelter date back to the terminal Pleistocene, but the burials are bracketed between circa 8200 and 9500BP. The collection contains only six skulls well-enough preserved to be measured. The Santana do Riacho late Paleoindians present a cranial morphology characterized by long and narrow neurocrania, low and narrow faces, with low nasal apertures and orbits. The multivariate analyses show that they exhibit strong morphological affinities with present day Australians and Africans, showing no resemblance to recent Northern Asians and Native Americans. These findings confirm our long held opinion that the settlement of the Americas was more complicated in terms of biological input than has been widely assumed. The working hypothesis is that two very distinct populations entered the New World by the end of the Pleistocene, and that the transition between the cranial morphology of the Paleoindians and the morphology of later Native Americans, which occurred around 8-9ka, was abrupt. This, in our opinion, is a more parsimonious explanation for the diversity detected than a long, local microevolutionary process mediated by selection and drift. The similarities of the first South Americans with sub-Saharan Africans may result from the fact that the non-Mongoloid Southeast Asian ancestral population came, ultimately, from Africa, with no major modification in the original cranial bau plan of the first modern humans.     

mtDNA control-region sequence variation suggests multiple independent origins of an "Asian-specific" 9-bp deletion in sub-Saharan Africans.

The intergenic COII/tRNA(Lys) 9-bp deletion in human mtDNA, which is found at varying frequencies in Asia, Southeast Asia, Polynesia, and the New World, was also found in 81 of 919 sub-Saharan Africans. Using mtDNA control-region sequence data from a subset of 41 individuals with the deletion, we identified 22 unique mtDNA types associated with the deletion in Africa. A comparison of the unique mtDNA types from sub-Saharan Africans and Asians with the 9-bp deletion revealed that sub-Saharan Africans and Asians have sequence profiles that differ in the locations and frequencies of variant sites. Both phylogenetic and mismatch-distribution analysis suggest that 9-bp deletion arose independently in sub-Saharan Africa and Asia and that the deletion has arisen more than once in Africa. Within Africa, the deletion was not found among Khoisan peoples and was rare to absent in western and southwestern African populations, but it did occur in Pygmy and Negroid populations from central Africa and in Malawi and southern African Bantu-speakers. The distribution of the 9-bp deletion in Africa suggests that the deletion could have arisen in central Africa and was then introduced to southern Africa via the recent "Bantu expansion."     

Dorset and Thule divergence from east Central Asian roots

The history of the immigration of East Asians to America during the last glacial period remains controversial. In an attempt to add critical data to this problem, a large sample of whole teeth derived from Southeast Asian, Mongolian, Thule, Western Inuit, and pre-Inca (Huari) people was quantified (N = 4,507 teeth from 495 individuals; ∼30 variables per tooth). Multivariate analysis helped establish that all Native Americans were likely derived from one ancient, extinct population that resided in the region of Mongolia (east Central Asia), and that Mongolians and Southeast Asians are two independent groups. A controversial and enigmatic Central Canadian Arctic “Thule culture Inuit” group on Southampton Island that survived until 1902 was identified as a relic, mainly Paleoeskimo Dorset community. Surprisingly, there was little, or no, indication of Dorset-to-Thule gene flow. Cumulatively, the data suggest that a small population of Paleoindian founders remained resident in Beringia, may have blocked further immigration, and were the antecedents to the Thule/Inuit. With the confluence of the Arctic and Pacific oceans at the breakup of Beringia, the resulting increased availability of marine animal food sources allowed this population to increase in size and expand throughout the eastern Arctic. Am J Phys Anthropol 106:207–218, 1998. © 1998 Wiley-Liss, Inc.     

Negritos, Australian aborigines, and the "proto-sundadont" dental pattern: the basic populations in east Asia, V.

Five evolutionarily significant dental traits were identified from a B-square distance analysis of nine crown characters recorded for several populations of East Asia and Oceania. Intergroup variation in these traits distinguishes three major divisions of the Mongoloid dental complex: sundadonty, sinodonty, and the dental pattern of Australian Aborigines. The Australian crown features may be characterized as having high frequencies of evolutionarily conservative characters. Negritos, one of the probable representatives of indigenous inhabitants of Southeast Asia who may have shared a common ancestor with Australians, possess the more derived sundadont dental pattern. As far as the five crown traits treated here are concerned, Australian dental features may be described as conforming to a "proto-sundadont" dental pattern, applying Turner's terminology. This pattern may represent a microevolutionary step prior to the emergence of the sundadont and sinodont patterns.