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 morphology of the Dawenkou Neolithic population in North China: implications for the origin and distribution of Sinodonty

We compare the incidence of 25 nonmetric dental traits of the people of the Neolithic Dawenkou culture (6300-4500 BP) sites in Shandong Province, North China with those of other East Asian populations. The Dawenkou teeth had an overwhelmingly greater resemblance to the Sinodont pattern typical of Northeast Asia than to the Sundadont pattern typical of Southeast Asia. Multidimensional scaling using Smith's mean measure of divergence (MMD) statistic place the Dawenkou sample near the Amur and the North China-Mongolia populations in the area of the plot indicating typical Sinodonty. The existence of the Sinodont population in Neolithic North China suggests a possible continuity of Sinodonty from the Upper Cave population at Zhoukoudian (about 34000-10000 BP) to the modern North Chinese. The presence of Sinodonty in Shandong Province shows that the Japan Sea and East China Sea were strong barriers to gene flow for at least 3000 years, because at this time the Jomonese of Japan were fully Sundadont. In addition, we suggest that the descendants of the Dawenkou population cannot be excluded as one of the source populations that contributed to sinodontification in Japan.     

Y chromosome genetic variation in the Italian peninsula is clinal and supports an admixture model for the Mesolithic-Neolithic encounter

The Italian peninsula, given its geographical location in the middle of the Mediterranean basin, was involved in the process of the peopling of Europe since the very beginning, with first settlements dating to the Upper Paleolithic. Later on, the Neolithic revolution left clear evidence in the archeological record, with findings going back to 7000 B.C. We have investigated the demographic consequences of the agriculture revolution in this area by genotyping Y chromosome markers for almost 700 individuals from 12 different regions. Data analysis showed a non-random distribution of the observed genetic variation, with more than 70% of the Y chromosome diversity distributed along a North-South axis. While the Greek colonisation during classical time appears to have left no significant contribution, the results support a male demic diffusion model, even if population replacement was not complete and the degree of Neolithic admixture with Mesolithic inhabitants was different in different areas of Italy.     

Paleolithic and neolithic lineages in the European mitochondrial gene pool.

Phylogenetic and diversity analysis of the mtDNA control region sequence variation of 821 individuals from Europe and the Middle East distinguishes five major lineage groups with different internal diversities and divergence times. Consideration of the diversities and geographic distribution of these groups within Europe and the Middle East leads to the conclusion that ancestors of the great majority of modern, extant lineages entered Europe during the Upper Paleolithic. A further set of lineages arrived from the Middle East much later, and their age and geographic distribution within Europe correlates well with archaeological evidence for two culturally and geographically distinct Neolithic colonization events that are associated with the spread of agriculture. It follows from this interpretation that the major extant lineages throughout Europe predate the Neolithic expansion and that the spread of agriculture was a substantially indigenous development accompanied by only a relatively minor component of contemporary Middle Eastern agriculturalists. There is no evidence of any surviving Neanderthal lineages among modern Europeans.     

A Comparison of Y-Chromosome Variation in Sardinia and Anatolia Is More Consistent with Cultural Rather than Demic Diffusion of Agriculture

Two alternative models have been proposed to explain the spread of agriculture in Europe during the Neolithic period. The demic diffusion model postulates the spreading of farmers from the Middle East along a Southeast to Northeast axis. Conversely, the cultural diffusion model assumes transmission of agricultural techniques without substantial movements of people. Support for the demic model derives largely from the observation of frequency gradients among some genetic variants, in particular haplogroups defined by single nucleotide polymorphisms (SNPs) in the Y-chromosome. A recent network analysis of the R-M269 Y chromosome lineage has purportedly corroborated Neolithic expansion from Anatolia, the site of diffusion of agriculture. However, the data are still controversial and the analyses so far performed are prone to a number of biases. In the present study we show that the addition of a single marker, DYSA7.2, dramatically changes the shape of the R-M269 network into a topology showing a clear Western-Eastern dichotomy not consistent with a radial diffusion of people from the Middle East. We have also assessed other Y-chromosome haplogroups proposed to be markers of the Neolithic diffusion of farmers and compared their intra-lineage variation—defined by short tandem repeats (STRs)—in Anatolia and in Sardinia, the only Western population where these lineages are present at appreciable frequencies and where there is substantial archaeological and genetic evidence of pre-Neolithic human occupation. The data indicate that Sardinia does not contain a subset of the variability present in Anatolia and that the shared variability between these populations is best explained by an earlier, pre-Neolithic dispersal of haplogroups from a common ancestral gene pool. Overall, these results are consistent with the cultural diffusion and do not support the demic model of agriculture diffusion.     

Who were the ancient Egyptians? Dental affinities among Neolithic through postdynastic peoples

Qualitative and quantitative methods are employed to describe and compare up to 36 dental morphological variants in 15 Neolithic through Roman-period Egyptian samples. Trait frequencies are determined, and phenetic affinities are calculated using the mean measure of divergence and Mahalanobis D2 statistics for discrete traits; the most important traits in generating this intersample variation are identified with correspondence analysis. Assuming that the samples are representative of the populations from which they derive, and that phenetic similarity provides an estimate of genetic relatedness, these affinities are suggestive of overall population continuity. That is, other than a few outliers exhibiting extreme frequencies of nine influential traits, the dental samples appear to be largely homogenous and can be characterized as having morphologically simple, mass-reduced teeth. These findings are contrasted with those resulting from previous skeletal and other studies, and are used to appraise the viability of five Egyptian peopling scenarios. Specifically, affinities among the 15 time-successive samples suggest that: 1) there may be a connection between Neolithic and subsequent predynastic Egyptians, 2) predynastic Badarian and Naqada peoples may be closely related, 3) the dynastic period is likely an indigenous continuation of the Naqada culture, 4) there is support for overall biological uniformity through the dynastic period, and 5) this uniformity may continue into postdynastic times.     

Concordance of cranial and dental morphological traits and evidence for endogamy in ancient Egypt

A biological affinities study based on frequencies of cranial nonmetric traits in skeletal samples from three cemeteries at predynastic Naqada, Egypt, confirms the results of a recent nonmetric dental morphological analysis. Both cranial and dental traits analyses indicate that the individuals buried in a cemetery characterized archaeologically as high status are significantly different from individuals buried in two other, apparently nonelite cemeteries and that the nonelite samples are not significantly different from each other. A comparison with neighbouring Nile Valley skeletal samples suggests that the high status cemetery represents an endogamous ruling or elite segment of the local population at Naqada, which is more closely related to populations in northern Nubia than to neighbouring populations in southern Egypt. © 1996 Wiley-Liss, Inc.     

A Predominantly Neolithic Origin for European Paternal Lineages

The relative contributions to modern European populations of Paleolithic hunter-gatherers and Neolithic farmers from the Near East have been intensely debated. Haplogroup R1b1b2 (R-M269) is the commonest European Y-chromosomal lineage, increasing in frequency from east to west, and carried by 110 million European men. Previous studies suggested a Paleolithic origin, but here we show that the geographical distribution of its microsatellite diversity is best explained by spread from a single source in the Near East via Anatolia during the Neolithic. Taken with evidence on the origins of other haplogroups, this indicates that most European Y chromosomes originate in the Neolithic expansion. This reinterpretation makes Europe a prime example of how technological and cultural change is linked with the expansion of a Y-chromosomal lineage, and the contrast of this pattern with that shown by maternally inherited mitochondrial DNA suggests a unique role for males in the transition.     

On the trail of Neolithic mice and men towards Transcaucasia: zooarchaeological clues from Nakhchivan (Azerbaijan)

Transcaucasia comprises a key region for understanding the history of both the hybrid zone between house mouse lineages and the dispersal of the Neolithic way of life outside its Near Eastern cradle. The opportunity to document the colonization history of both men and mice in Transcaucasia was made possible by the discovery of mouse remains accumulated in pits from a 6000‐year‐old farming village in the Nakhchivan (Autonomous Republic of Nakhchivan, Azerbaijan). The present study investigated their taxonomy and most likely dispersal path through the identification of the Mus lineage to which they might belong using a geometric morphometric approach of dental traits distances between archaeological and modern populations of the different Mus lineages of South‐West Asia. We demonstrate that the mouse remains trapped in the deep storage pits of the dwelling belong to the Mus musculus domesticus from the Near East, with dental shapes similar to current populations in Northern Syria. These results strongly suggest that the domesticus lineage was dispersed into Transcaucasia from the upper Euphrates valley by Neolithic migration, some time between the 7th and 5th millennium BC, providing substantial evidence to back up the scenario featuring near‐eastern stimuli in the emergence of agriculture in the South Caucasus. The domesticus mitochondrial DNA signature of the current house mouse in the same location 5000 years later, as well as their turnover towards a subspecies musculus/castaneus phenotype, suggests that early domesticus colonizers hybridized with a later musculus (and maybe castaneus) dispersal originating from south of the Caspian Sea and/or Northern Caucasia. © 2013 The Linnean Society of London     

Population continuity vs. discontinuity revisited: dental affinities among late Paleolithic through Christian-era Nubians

The present study revisits a subject that has been a source of long-standing bioarchaeological contention, namely, estimation of Nubian population origins and affinities. Using the Arizona State University dental anthropology system, frequencies of 36 crown, root, and intraoral osseous discrete traits in 12 late Pleistocene through early historic Nubian samples were recorded and analyzed. Specifically, intersample phenetic affinities, and an indication of which traits are most important in driving this variation, were determined through the application of correspondence analysis and the mean measure of divergence distance statistic. The results support previous work by the author and others indicating that population discontinuity, in the form of replacement or significant gene flow into an existing gene pool, occurred sometime after the Pleistocene. This analysis now suggests that the break occurred before the Final Neolithic. Samples from the latter through Christian periods exhibit relative homogeneity, which implies overall post-Pleistocene diachronic and regional population continuity. Yet there are several perceptible trends among these latter samples that: 1) are consistent with documented Nubian population history, 2) enable the testing of several existing peopling hypotheses, and 3) allow the formulation of new hypotheses, including a suggestion of two post-Pleistocene subgroups predicated on an age-based sample dichotomy.     

Y-chromosomal diversity in Europe is clinal and influenced primarily by geography, rather than by language

Clinal patterns of autosomal genetic diversity within Europe have been interpreted in previous studies in terms of a Neolithic demic diffusion model for the spread of agriculture; in contrast, studies using mtDNA have traced many founding lineages to the Paleolithic and have not shown strongly clinal variation. We have used 11 human Y-chromosomal biallelic polymorphisms, defining 10 haplogroups, to analyze a sample of 3,616 Y chromosomes belonging to 47 European and circum-European populations. Patterns of geographic differentiation are highly nonrandom, and, when they are assessed using spatial autocorrelation analysis, they show significant clines for five of six haplogroups analyzed. Clines for two haplogroups, representing 45% of the chromosomes, are continentwide and consistent with the demic diffusion hypothesis. Clines for three other haplogroups each have different foci and are more regionally restricted and are likely to reflect distinct population movements, including one from north of the Black Sea. Principal-components analysis suggests that populations are related primarily on the basis of geography, rather than on the basis of linguistic affinity. This is confirmed in Mantel tests, which show a strong and highly significant partial correlation between genetics and geography but a low, nonsignificant partial correlation between genetics and language. Genetic-barrier analysis also indicates the primacy of geography in the shaping of patterns of variation. These patterns retain a strong signal of expansion from the Near East but also suggest that the demographic history of Europe has been complex and influenced by other major population movements, as well as by linguistic and geographic heterogeneities and the effects of drift.     

Population Genomic Analysis of Ancient and Modern Genomes Yields New Insights into the Genetic Ancestry of the Tyrolean Iceman and the Genetic Structure of Europe

Genome sequencing of the 5,300-year-old mummy of the Tyrolean Iceman, found in 1991 on a glacier near the border of Italy and Austria, has yielded new insights into his origin and relationship to modern European populations. A key finding of that study was an apparent recent common ancestry with individuals from Sardinia, based largely on the Y chromosome haplogroup and common autosomal SNP variation. Here, we compiled and analyzed genomic datasets from both modern and ancient Europeans, including genome sequence data from over 400 Sardinians and two ancient Thracians from Bulgaria, to investigate this result in greater detail and determine its implications for the genetic structure of Neolithic Europe. Using whole-genome sequencing data, we confirm that the Iceman is, indeed, most closely related to Sardinians. Furthermore, we show that this relationship extends to other individuals from cultural contexts associated with the spread of agriculture during the Neolithic transition, in contrast to individuals from a hunter-gatherer context. We hypothesize that this genetic affinity of ancient samples from different parts of Europe with Sardinians represents a common genetic component that was geographically widespread across Europe during the Neolithic, likely related to migrations and population expansions associated with the spread of agriculture.     

Neolithic transition and dental changes: the case of an Italian site

This paper compares the dentition of two samples of populations which lived in the same sites of Western Liguria (Italy) during the late phases of a hunting and gathering economy and at the beginning of agriculture respectively. The characteristics considered include metrics, pathology and wear. The older sample dates back to the Late Epigravettian (X-IX millennium B.C.), while the group of early farmers belongs mostly to the Middle Neolithic (IV millennium B.C.).The Neolithic group, conforming to the trend generally observed in human populations after the development of agriculture, shows reduction in crown size, spread of caries, increase in enamel hypoplasia, slower rates of wear on posterior teeth and less use of the anterior part of the dentition. Economic, technological, and dietary differences can provide a coherent explanatory model for the observed results. The possibility of biological relationships between the two samples is discussed with reference to archeological data.     

Assessing change in diet and biological affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura: A preliminary dental comparison of Feteira II and Bolores

Although the social and political changes accompanying the transition from the Neolithic through Copper Age, between the 4th and 3rd millennia cal BCE, in southwestern Iberia are reasonably well understood, much less is known about whether population movements and dietary changes accompanied these transformations. To address this question, human dental remains from the Middle through Late Neolithic site of Feteira II (3600–2900 cal BCE) and the Late Neolithic site of Bolores (2800–2600 cal BCE) in the Portuguese Estremadura were used to examine diet (microwear) and affinity (dental non-metrics). Microwear features were not found to be significantly different between Feteira II and Bolores, suggesting that the emergence of social complexity during this period did not result in large-scale changes in subsistence practices during the period of use at these sites. Using the Arizona State University Dental Anthropology System and supporting statistics, no significant difference between the samples from Feteira II and Bolores was observed, suggesting that no population replacement occurred between the Middle Neolithic and Late Neolithic/Copper Age. However, at Bolores there is some indication that there may have been demographic exchanges between southern Iberian and North African populations during the Late Neolithic/Copper Age.     

Implications of Dental Caries in Anatolia: From Hunting–Gathering to the Present

Teeth are considered one of the most informative and durable parts of the skeleton. In Dental Anthropology, they are used to obtain information on culture, health, diet, variability and evolutionary trends in dental morphology as well as development, eruption and dental pathologies in the past and modern populations. In this study, an attempt was made to evaluate the results of dental caries, which is the most common dental disease, in order to document changing patterns of health and diet ranging from the transitional period of hunting–gathering through agriculture to the present day in human history in Anatolia. From the total sample of 400 modern individuals, a total of 5,208 maxillary teeth and 5,153 mandibular teeth were studied. The percentage of the occurrence of dental caries based on the individuals was 77.8%, whereas the frequency of dental caries on tooth type and class was 17.1% (18.0% maxillary decay; 16.2% mandibular decay). A comparative study of the frequency of caries in certain periods indicates the following: in the hunting–gathering period it was 1%–2%, in the Early Neolithic it was 3%–5% (Catal Hoyuk), in the Neolithic (beginning of agriculture) it was 5.6% (Cayonu), in the Chalcolithic it was 11.7% (Norsuntepe), in the Roman period it was 11.1% (Panaztepe), and 16% (Datca), in the Late Byzantine it was 10.9% (Iznik) and in the Medieval it is 14.2% (Arslantepe). These findings contribute to understanding how dietary change and life conditions are interrelated with the changing patterns of dental diseases in Anatolian populations.     

Tooth wear and dental pathology at the advent of agriculture: new evidence from the Levant

Differences in patterns of diet and subsistence through the analysis of dental pathology and tooth wear were studied in skeletal populations of Natufian hunter-gatherers (10,500-8300 BC) and Neolithic populations (8300-5500 BC, noncalibrated) from the southern Levant. 1,160 Natufians and 804 Neolithic teeth were examined for rate of attrition, caries, antemortem tooth loss, calculus, periapical lesions, and periodontal processes. While the Natufian people manifest a higher rate of dental attrition and periodontal disease (36.4% vs. 19%), Neolithic people show a higher rate of calculus. Both populations manifested low and similar rates of caries (6.4% in the Natufian vs. 6.7% in the Neolithic), periapical lesions (not over 1.5%), and antemortem tooth loss (3.7% vs. 4.5%, respectively). Molar wear pattern in the Neolithic is different than in the Natufian. The current study shows that the dental picture obtained from the two populations is multifactorial in nature, and not exclusively of dietary origin, i.e., the higher rate and unique pattern of attrition seen in the Natufian could result from a greater consumption of fibrous plants, the use of pestles and mortars (which introduce large quantities of stone-dust to the food), and/or the use of teeth as a "third hand." The two major conclusions of this study are: 1) The transition from hunting and gathering to a food-producing economy in the Levant did not promote changes in dental health, as previously believed. This generally indicates that the Natufians and Neolithic people of the Levant may have differed in their ecosystem management (i.e., gathering vs. growing grains), but not in the type of food consumed. 2) Changes in food-preparation techniques and nondietary usage of the teeth explain much of the variation in tooth condition in populations before and after the agricultural revolution.     

Genomic affinities of two 7,000-year-old iberian hunter-gatherers

The genetic background of the European Mesolithic and the extent of population replacement during the Neolithic [1-10] is poorly understood, both due to the scarcity of human remains from that period [11-18] and the inherent methodological difficulties of ancient DNA research. However, advances in sequencing technologies are both increasing data yields and providing supporting evidence for data authenticity, such as nucleotide misincorporation patterns [19-22]. We use these methods to characterize both the mitochondrial DNA genome and generate shotgun genomic data from two exceptionally well-preserved 7,000-year-old Mesolithic individuals from La Bra?a-Arintero site in León (Northwestern Spain) [23]. The mitochondria of both individuals are assigned to U5b2c1, a haplotype common among the small number of other previously studied Mesolithic individuals from Northern and Central Europe. This suggests a remarkable genetic uniformity and little phylogeographic structure over a large geographic area of the pre-Neolithic populations. Using Approximate Bayesian Computation, a model of genetic continuity from Mesolithic to Neolithic populations is poorly supported. Furthermore, analyses of 1.34% and 0.53% of their nuclear genomes, containing about 50,000 and 20,000 ancestry informative SNPs, respectively, show that these two Mesolithic individuals are not related to current populations from either the Iberian Peninsula or Southern Europe.     

Complete mitochondrial genomes reveal neolithic expansion into Europe

The Neolithic transition from hunting and gathering to farming and cattle breeding marks one of the most drastic cultural changes in European prehistory. Short stretches of ancient mitochondrial DNA (mtDNA) from skeletons of pre-Neolithic hunter-gatherers as well as early Neolithic farmers support the demic diffusion model where a migration of early farmers from the Near East and a replacement of pre-Neolithic hunter-gatherers are largely responsible for cultural innovation and changes in subsistence strategies during the Neolithic revolution in Europe. In order to test if a signal of population expansion is still present in modern European mitochondrial DNA, we analyzed a comprehensive dataset of 1,151 complete mtDNAs from present-day Europeans. Relying upon ancient DNA data from previous investigations, we identified mtDNA haplogroups that are typical for early farmers and hunter-gatherers, namely H and U respectively. Bayesian skyline coalescence estimates were then used on subsets of complete mtDNAs from modern populations to look for signals of past population expansions. Our analyses revealed a population expansion between 15,000 and 10,000 years before present (YBP) in mtDNAs typical for hunters and gatherers, with a decline between 10,000 and 5,000 YBP. These corresponded to an analogous population increase approximately 9,000 YBP for mtDNAs typical of early farmers. The observed changes over time suggest that the spread of agriculture in Europe involved the expansion of farming populations into Europe followed by the eventual assimilation of resident hunter-gatherers. Our data show that contemporary mtDNA datasets can be used to study ancient population history if only limited ancient genetic data is available.     

郑州青台遗址新石器时代中晚期人群龋齿的统计与分析

本文通过对郑州青台遗址新石器时代中晚期91例个体、1913枚牙齿罹患龋齿的统计与分析可知,青台人群患龋率为71.43%,龋齿率为13.38%。其中,龋齿率女性高于男性,可能与女性孕期生理变化、食物选择及性别分工等有关。上颌龋齿率高于下颌,臼齿及咬合面为龋齿易患齿类及部位。通过对比可知青台人群显示出较高的龋齿罹患率,暗示该人群饮食中应包含较多的碳水化合物类食物,这可能与新石器时代中晚期黄河中游发达的旱作农业有关。此外,龋齿率在黄河及长江中、下游新石器时代农业人群中的区域性差异可能与龋病病因的复杂性和各地区不同的文化面貌、人群生活方式有关。     

Prehistoric "fish-eaters" along the eastern Arabian coasts: dental variation, morphology, and oral health in the Ra's al-Hamra community (Qurum, sultanate of Oman, 5th-4th millennia BC)

The Ra's al-Hamra prehistoric fishermen lived in isolation on the Qurum rocky promontorium in Oman during the 5th-4th millennia BC. To date, they represent the most ancient and numerous human fossil group excavated from the Arabian peninsula. Like other contemporaneous archaeologically documented small communities along the desert Arabian coasts, they intensively exploited ocean resources and collected molluscs from nearby mangrove swamps. The present study analyzes aspects of dental anthropology (including crown variation, morphology, dental wear, and oral health), in 600 permanent teeth from 49 individuals of both sexes excavated at the Mesolithic RH5-site by the Italian Archaeological Mission in Oman from 1981 to 1985. In association with a general low degree of morphometric variation, the Ra's al-Hamra dental crowns show low sexual dimorphism and are consistently reduced in size. These features are unexpected in a preagricultural population, especially when these data are compared to other eastern African and near Middle Eastern prehistoric populations. These data are discussed within the general context of human dental structural reduction occurring during the post-Pleistocene and are interpreted according to the "increasing population density effect" model. There are other significant differences that characterize the Ra's al-Hamra dentitions with respect to both eastern and western prehistoric human groups. The frequency of numerous nonmetric crown traits supports the hypothesis that a microdifferentiation phenomenon occurred in this marginal area. The preliminary skeletal analysis and the palaeodemographic profile show that the Omani prehistoric fishermen were affected by genetic isolation and inbreeding as well as strong environmental stress. Because of the grit assimilated with dried fish and the high shellfish consumption, dental wear was extreme in all age groups at Ra's al-Hamra and occasionally was responsible for serious hematogenously spread infections. In spite of the great anthropological importance of ancient Arabian populations, very few studies on skeletal and dental samples have been completed. The present paper offers an odontological data set for future comparative research in the area.