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.     

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.     

Canary islands-north African population affinities: measures of divergence based on dental morphology

This investigation addresses two related questions about the origins and biological affinities of the Canary Islands' aboriginal inhabitants. First: With which North African populations do the pre-conquest inhabitants of the Canary Islands have their greatest affinities? Second: Does inter-island biological variability among the Canary Islanders, as has been suggested by other researchers (Hooton 1925, Schwidetzky 1963), imply that potentially different founding populations remained distinct during the pre-conquest period? This study employs dental morphology data derived from pre-conquest skeletons to answer these questions. Non-metric dental traits appear to be controlled by polygenic systems with a low to moderate environmental contribution to the resulting phenotype (Berry 1978, Harris & Bailit 1980, Nichol 1990) and can thus be assumed to reflect genetic relationships. The dental morphology of a sample of Canary Islanders (n = 397) is compared to that of Northwest African samples of Algerian Shawia Berbers (n = 26), Kabyle Berbers (n = 32), Bedouin Arabs (n = 49) and Punic Carthaginians (n = 28) as well as to six samples from Northeast Africa (n = 307) included for the purpose of understanding Canary Islanders' affinities within a wider context. The analysis employs 28 dental traits, quantifying differences in their expression among the various samples through a summary statistic, CAB Smith's Mean Measure of Divergence (MMD). The MMD analysis indicates that the Canary Island sample is most similar to the four samples from Northwest Africa: the Shawia Berbers, Kabyle Berbers, Bedouin Arabs and Carthaginians, less similar to the three Egyptian samples and least like the three Nubian samples. An intra-island comparison among samples from La Gomera, Gran Canaria and Tenerife reveals low, insignificant MMD values in all cases, implying that inter-island dental morphology differences are not so great as to require hypotheses of separate founding populations.     

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.     

Origins and spread of agriculture in Italy: a nonmetric dental analysis

Dental morphological traits were employed in this study as direct indicators of biological affinities among the populations that inhabited the Italian peninsula from the Upper Paleolithic-Mesolithic to Medieval times. Our analysis aims at contributing to the ongoing debate regarding the origin and spread of agriculture in the peninsula by contrasting the dental evidence of archaeological and modern molecular samples. It is not possible to generalize given the complex and dynamic nature of these populations. However, the results from the principal component analysis, maximum likelihood, mean measure of divergence, and multidimensional scaling do indicate a net separation of the Paleo-Mesolithic sample from the other groups that is not related to dental reduction. This suggests that the shift in dental morphology was the product of Neolithic populations migrating into the peninsula from other areas. Nonetheless, the Paleo-Mesolithic populations share several discriminative traits with the Neolithic group. The biological relevance of such evidence suggests that, to some minor extent, the spread of agriculture did not occur by total population replacement. Because of regional small sample sizes, this hypothesis cannot be tested on a micro-regional scale. It is, however, feasible to depict a scenario where processes of genetic mixture or replacement probably took place at different rates on a macro-regional level.     

Nonmetric tooth crown traits in a Sri Lankan aboriginal Vedda population

This study was conducted to determine the frequencies of non-metric tooth crown traits of Vedda of Sri Lanka and to investigate the affinities of these morphological variations with those of other world populations.Fifty dental plaster casts were observed. The Arizona State University dental anthropology system was adopted for classification of the 16 traits observed. We used 13 traits to compare the Vedda and other world populations. Using the frequencies of 13 traits, Smith Mean Measure of Divergence was calculated to determine inter-population distances. Affinities among the Vedda and other world populations were expressed in two dimensions of the principal coordinate analysis.Cusp number in mandibular second molar and hypocone absence in maxillary second molar had the highest frequency at 95.9% and 93.8%, respectively. Shovelling, double shovelling in the maxillary central incisor and deflecting wrinkle in the mandibular first molar had the lowest frequency at 0%. The principal coordinate analysis showed that Sino American and Western Eurasian populations were separated in negative and positive directions in the first principal coordinate axis. Vedda located with the Western Eurasian population groups. Sahul and Sunda Pacific populations located in the intermediate position between Sino American and Western Eurasian populations.The dental phenotype of Vedda has close affinities with those of early south Asian populations. They are far different from Sino American and Sunda pacific populations. Vedda shows closer affinities to Sahul Pacific and South African (Bantu) populations.     

Cranial variation in the terminal Late Archaic of Ohio

Cranial and mandibular discrete traits and cranial metric traits were collected from 99-125 individuals in eight terminal Late Archaic sites. The analysis of ten metric traits in six samples showed that the samples shared the same generalized variance and that sexual dimorphism in the means of the metric traits was greater than inter-site differences. Since these samples share the same size and shape expression of a complex set of polygenic traits, we hypothesize a historical relationship among these samples. Discrete trait analysis showed a pattern of differentiation among the eight samples. This pattern of differentiation is related directly to the geographical separation between samples, and, as with metric traits, cultural differences do not contribute to the pattern of biological differences. The overall pattern of osteological variation in these samples can be summarized parsimoniously by paraphrasing the first law of geography: All of the terminal Late Archaic populations of Ohio were related to each other, but closer ones were more related than distant ones.     

Relationships between non-metric skeletal traits and cranial size and shape.

A long-standing controversy exists about the comparative utility of metric and non-metric traits as biological indicators in population studies. We hypothesize that the underlying scale which determines the presence or absence of a cranial non-metric trait is an expression of general and/or local size variation in the cranium. Therefore metric and non-metric traits will share a common developmental determination. The hypothesis implies that the underlying scale of a non-metric trait will be correlated with measures of cranial size and shape. Forty-eight cranial metric and twenty-five cranial non-metric traits were scored on the left side of adult male crania from four North American Indian populations. New threshold traits were generated for each non-metric trait by dichotomizing discriminant scores produced by discriminant function analysis. The discriminant analysis was performed using metric traits to discriminate between groups formed by non-metric trait presence or absence. Every non-metric trait tested was significantly correlated with its threshold trait in at least one population. The correlations were of moderate to high levels depending on the trait and population sample studied. This implies that metric and non-metric traits share a moderate to high degree of developmental determination. The cause of these correlations may lie in the common effects that growth and development of the soft tissue and functional spaces of the cranium exert on both metric and non-metric traits.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques of Cayo Santiago. III. Relative heritability of skeletal nonmetric and metric traits

This study addresses the long-standing controversy in skeletal biology concerning the relative utility of skeletal metric and nonmetric traits for studies of biological relationship. This controversy centers on the relative heritability of these two trait sets. This paper presents heritabilities for a series of skeletal metric and nonmetric traits measured with the same sample of mother-offspring pairs from the Cayo Santiago skeletal collection of rhesus macaques. Skeletal nonmetric traits display significantly greater heritability estimates than metric traits. This difference is due primarily to the high heritability estimates of hyperostotic nonmetric traits. Foraminal traits are not significantly more heritable than skeletal metric traits. The generality of this pattern of heritability values, in which hyperostotic nonmetric traits are more highly heritable than foraminal nonmetric and metric traits, depends on future empirical study of the correlation of heritability values in populations and theoretical work.     

Floral integration,phenotypic covariance structure and pollinator variation in bumblebee‐pollinated Helleborus foetidus

By analysing patterns of phenotypic integration and multivariate covariance structure of five metric floral traits in nine Iberian populations of bumblebee‐pollinated Helleborus foetidus (Ranunculaceae), this paper attempts to test the general hypothesis that pollinators enhance floral integration and selectively modify phenotypic correlations between functionally linked floral traits. The five floral traits examined exhibited significant phenotypic integration at all populations, and both the magnitude and the pattern of integration differed widely among populations. Variation in extent and pattern of integration was neither distance‐dependent nor significantly related to between‐population variation in taxonomical composition and morphological diversity of the pollinator assemblage. Patterns of floral integration were closer to expectations derived from consideration of developmental affinities between floral whorls than to expectations based on a pollinator‐mediated adaptive hypothesis. Taken together, results of this study suggest that between‐population differences in magnitude and pattern of floral integration in H. foetidus are probably best explained as a consequence of random genetic sampling in the characteristically small and ephemeral populations of this species, rather than reflecting the selective action of current pollinators.     

Ancient teeth and modern human origins: an expanded comparison of African Plio-Pleistocene and recent world dental samples

Previous research by the first author revealed that, relative to other modern peoples, sub-Saharan Africans exhibit the highest frequencies of ancestral (or plesiomorphic) dental traits and, thus, appear to be least derived dentally from an ancestral hominin state. This determination, in conjunction with various other lines of dental morphological evidence, was interpreted to be supportive of an African origin for modern humans. The present investigation expands upon this work by using: 1) direct observations of fossil hominin teeth, rather than data gleaned from published sources, 2) a single morphological scoring system (the Arizona State University Dental Anthropology System) with consistent trait breakpoints, and 3) data from larger and more varied modern human comparative samples. As before, a multivariate distance statistic, the mean measure of divergence, was used to assess diachronic phenetic affinities among the Plio-Pleistocene hominins and modern humans. The present study also employed principal components analysis on dental trait frequencies across samples. Both methods yielded similar results, which support the previous findings; that is, of all modern human samples, sub-Saharan Africans again exhibit the closest phenetic similarity to various African Plio-Pleistocene hominins-through their shared prevalence of morphologically complex crown and root traits. The fact that sub-Saharan Africans express these apparently plesiomorphic characters, along with additional information on their affinity to other modern populations, evident intra-population heterogeneity, and a world-wide dental cline emanating from the sub-continent, provides further evidence that is consistent with an African origin model.     

Questions of khoesan continuity: Dental affinities among the indigenous holocene peoples of South Africa

The present report follows up on the findings of previous research, including recent bioarchaeological study of well‐dated Khoesan skeletal remains, that posits long term biological continuity among the indigenous peoples of South Africa after the Pleistocene. The Arizona State University Dental Anthropology System was used to record key crown, root, and intraoral osseous nonmetric traits in six early‐through‐late Holocene samples from the Cape coasts. Based on these data, phenetic affinities and an identification of traits most important in driving intersample variation were determined using principal components analysis and the mean measure of divergence distance statistic. To expand biological affinity comparisons into more recent times, and thus preliminarily assess the dental impact of disproportionate non‐Khoesan gene flow into local peoples, dental data from historic Khoekhoe and San were also included. Results from the prehistoric comparisons are supportive of population continuity, though a sample from Matjes River Rockshelter exhibits slight phenetic distance from other early samples. This and some insignificant regional divergence among these coastal samples may be related to environmental and cultural factors that drove low‐level reproductive isolation. Finally, a close affinity of historic San to all samples, and a significant difference of Khoekhoe from most early samples is reflective of documented population history following immigration of Bantu‐speakers and, later, Europeans into South Africa. Am J Phys Anthropol 155:33–44, 2014. © 2014 Wiley Periodicals, Inc.     

Dental variation among four prehispanic Mexican populations

In this paper, the dental morphology of prehispanic Mesoamerican populations is described, compared, and examined within the context of New World dental variation. Twenty-eight morphological dental traits were studied and compared in four samples of prehispanic Mexican populations. After eliminating intra- and interobserver error, the dental morphological characteristics observed show evidence of heterogeneity among the populations. In particular, the oldest population, Tlatilco (1300–800 BC ), was significantly different from the other three groups, Cuicuilco (800–100 BC ), Monte Albán (500 BC –700 AD ) and Cholula (550–750 AD ). When the four samples were compared to other Mongoloid populations, either univariately or multivariately, it was observed that the Mexican groups did not follow a strict Sinodont (characteristic of Northeast Asia)/Sundadont (characteristic of Southeast Asia) classification (Turner [1979] Am. J. Phys. Anthropol. 51:619–636). From the traits examined, 27% presented frequencies consistent with Sinodont variation, while 73% of the traits showed similar incidence to Southeast Asian groups. Multivariately, the Mexican populations were found to fit an overall Sundadont classification. These results indicate that there is more dental morphological variation among American Indian populations than previously shown. © 1996 Wiley-Liss, Inc.     

Biological Affinities Between Migrant and Parental Populations in Punjab, India

In the present study, we have examined biological affinities between parental population, migrants from Pakistan after 1947 and non-Punjabi migrants from other states of India in Punjab. About 500 adult individuals of both sexes from 300 families among three groups were measured for 16 anthropometric and physiometric variables. Analysis of both univariate and multivariate suggests an overall significant migration effect in almost all variables especially anthropometric measurements between both sexes. The higher discrimination has been found in migrants from other states of India with reference to anthropometric traits as compared to other two populations. This is because of more genetic heterogeneity as would be expected from many states and many ethnicities of migrants. It was also found that the other two populations are more or less qualitatively similar, which reflects common ethnohistorical or geographic affiliations. However, overall, women showed a greater migration effect than men.     

Genetic Markers and Quantitative Genetic Variation in Medicago Truncatula (Leguminosae): A Comparative Analysis of Population Structure

Two populations of the selfing annual Medicago truncatula Gaertn. (Leguminoseae), each subdivided into three subpopulations, were studied for both metric traits (quantitative characters) and genetic markers (random amplified polymorphic DNA and one morphological, single-locus marker). Hierarchical analyses of variance components show that (1) populations are more differentiated for quantitative characters than for marker loci, (2) the contribution of both within and among subpopulations components of variance to overall genetic variance of these characters is reduced as compared to markers, and (3) at the population level, within population structure is slightly but not significantly larger for markers than for quantitative traits. Under the hypothesis that most markers are neutral, such comparisons may be used to make hypotheses about the strength and heterogeneity of natural selection in the face of genetic drift and gene flow. We thus suggest that in these populations, quantitative characters are under strong divergent selection among populations, and that gene flow is restricted among populations and subpopulations.     

The Iberomaurusian enigma: north African progenitor or dead end?

Data obtained during an ongoing dental investigation of African populations address two long-standing, hotly debated questions. First, was there genetic continuity between Late Pleistocene Iberomaurusians and later northwest Africans (e.g., Capsians, Berbers, Guanche)? Second, were skeletally-robust Iberomaurusians and northeast African Nubians variants of the same population? Iberomaurusians from Taforalt in Morocco and Afalou-Bou-Rhummel in Algeria, Nubians from Jebel Sahaba in Sudan, post-Pleistocene Capsians from Algeria and Tunisia, and a series of other samples were statistically compared using 29 discrete dental traits to help estimate diachronic local and regional affinities. Results revealed: (1) a relationship between the Iberomaurusians, particularly those from Taforalt, and later Maghreb and other North African samples, and (2) a divergence among contemporaneous Iberomaurusians and Nubian samples. Thus, some measure of long-term population continuity in the Maghreb and surrounding region is supported, whereas greater North African population heterogenity during the Late Pleistocene is implied.     

The phenetic distances of the living Druze from other human populations suggest a major genetic drift from the Western Eurasian ancestral category

The objectives were to determine the expression frequency and sexual dimorphism of 16 non-metric crown traits on the sample of permanent dentitions of the living Druze population (a Near Eastern genetic isolate) in Jordan, and to assess the biological affinity of this sample to 21 regional groups, and to the living general Jordanian population, based on these traits. Druze schoolchildren (46 males, 40 females; mean age = 16.0, sd = 0.5 years) were studied in 2011. The traits were classified using the Arizona State University dental anthropology system, counted with the individual count method, and dichotomized according to the criteria of Scott and Turner for the purpose of group comparisons. Fisher's exact test for dichotomized scores was used to assess sexual dimorphism in these traits. Smith's mean measure of divergence was used to measure all pairwise distance values among the groups. Sexual dimorphism was found in five traits (i.e., UI2 interruption grooves, 3-cusped UM2, UM1 Carabelli's tubercle/cusp, 4-cusp LM1, and LM2 Y-groove pattern). This study revealed that the dental pattern of living Druze, which is similar to that of the general Jordanian population, is sufficiently distinct from the Western Eurasian pattern and all other known dental patterns to form a distinct dental pattern for the regional group or subcategory to which these two populations belong. Moreover, the relatively large distance values of the living Druze and Jordanians from the other world groups considered, including the Western Eurasian groups, suggest a similar major genetic difference of these two populations from the Western Eurasian ancestry.     

Sahara: Barrier or corridor? Nonmetric cranial traits and biological affinities of North African late holocene populations

The Garamantes flourished in southwestern Libya, in the core of the Sahara Desert ~3,000 years ago and largely controlled trans-Saharan trade. Their biological affinities to other North African populations, including the Egyptian, Algerian, Tunisian and Sudanese, roughly contemporary to them, are examined by means of cranial nonmetric traits using the Mean Measure of Divergence and Mahalanobis D(2) distance. The aim is to shed light on the extent to which the Sahara Desert inhibited extensive population movements and gene flow. Our results show that the Garamantes possess distant affinities to their neighbors. This relationship may be due to the Central Sahara forming a barrier among groups, despite the archaeological evidence for extended networks of contact. The role of the Sahara as a barrier is further corroborated by the significant correlation between the Mahalanobis D(2) distance and geographic distance between the Garamantes and the other populations under study. In contrast, no clear pattern was observed when all North African populations were examined, indicating that there was no uniform gene flow in the region.     

Asymmetry and diversity of dermatoglyphic traits: population comparison in five endogamous groups of West Bengal, India

Five different endogamous populations encompassing the main social ranks in the caste hierarchy of West Bengal, India were analyzed. To compare variability in populations with contrasting ethnohistorical backgrounds, analysis of variance, Scheffe's test and cluster analysis were performed, as based on dermatoglyphic variables, namely, 22 quantitative traits and 36 indices of diversity and asymmetry. The present study reveals that: 1. Overall disparities among the 5 populations are expressed only in finger ridge counts on the Ist and Vth digits and PII, in a-b ridge counts, in endings of main lines A and D, and in MLI on the palms; 2. Heterogeneity is greater in fluctuating asymmetry than in directional asymmetry; 3. There is a greater heterogeneity in the 22 quantitative traits than in the 36 indices of diversity and asymmetry, with females contributing more than the males; 4. The highest contribution to population variation is by Lodha among five populations; 5. Inter-group variations are homogeneous in most of the variables, which does not correspond with the relationships to caste hierarchy of these populations; 6. The dendrograms based on dermatoglyphic variables demonstrate that the traditional grouping of Indian populations, based on caste hierarchy, may not be a reflection of their genetic origin, in that the pattern of clustering corresponded best with the known ethnohistorical records of the studied populations; 7. Hence, dermatoglyphic affinities may prove quite useful in tracing the ethnohistorical background of populations.     

红松种子园种群表型多样性研究

为揭示红松（Pinus koraiensis Sieb.et Zucc.）不同种群的表型分化程度和变异规律,以吉林省露水河红松种子园6个种群红松为研究对象,采用巢式方差分析、多重比较、变异系数和聚类分析等方法对红松的叶片、球果和种子共15个表型性状进行了系统分析。结果表明：（1）红松15个表型性状在种群间和种群内均存在着极显著的差异,红松种群遗变异比较丰富,在松属植物中属于中等水平,其中纬度最低的露水河种群在其中10个表型性状均值上表现出最高值;（2）红松种群间的表型分化系数（Vst）均值为12.39%,种群内的变异（87.61%）大于种群间的变异（12.39%）,种群内的变异是主要变异来源;（3）各表型性状平均变异系数为13.28%,变幅为6.16%-31.48%,叶片、球果、种子的平均变异系数依次为：球果17.86% > 针叶11.19% > 种子9.87%,种子性状最小,成为最稳定的表型性状,带岭和丰林种群表型性状遗传多样性要高于其他种群;（4）利用欧氏平均距离对红松种群进行UPGMA聚类分析,红松种群的表型性状按照地理距离而聚类,可将红松种子园6个种群分为4类,其表型性状跟地形（东北山脉）有一定的契合。红松种群具有中等水平的表型遗传多样性,种群间和种群内均存在丰富的表型变异,研究结果为顺利开展红松种质资源收集、保存,以及良种选育等工作提供依据。