Deciduous dental morphology of the prehistoric Jomon people of Japan: Comparison of nonmetric characters

Morphological variations of the deciduous dentition are as useful as those of the permanent dentition for determining the biological affinities of human populations. This paper provides material on morphological variations of deciduous teeth of the prehistoric Japanese population from the Late and the Latest Jomon Period (ca. 2000–ca. 300 B.C.). The expression of nonmetric traits of the deciduous teeth in the Jomon sample shows a closer affinity with modern Japanese and Native American samples than with American White, Asiatic Indian, and African samples. However, the frequency of shoveling in deciduous upper incisors in the Jomon sample is lower than those in modern Japanese and Native American samples. The Jomon sample also expresses a much higher frequency of cusp 6 in deciduous lower second molars than seen in modern Japanese, Ainu, and Native American samples. The frequency in the Jomon sample is equal to that in the Australian Aboriginal sample, which shows cusp 6 most frequently among the samples compared. A somewhat low incidence of incisor shoveling in the Jomon sample was also reported in the permanent dentition (Turner [1976] Science 193:911–913, [1979] Am. J. Phys. Anthropol. 51:619–635, [1987] Am. J. Phys. Anthropol. 73:305–321, [1990] Am. J. Phys. Anthropol. 82:295–317; T. Hanihara [1992] Am. J. Phys. Anthropol. 88:163–182, 88:183–196). However, the frequency of cusp 6 in the Jomon sample shows no significant difference from those of Northeast Asian or Native American samples in the permanent dentition (Turner [1987] Am. J. Phys. Anthropol. 73:305–321; T. Hanihara [1992] Am. J. Phys. Anthropol. 88:1–182, 88:183–196). Evidently, some nonmetric traits express an inter-group difference only in the deciduous dentition. © 1995 Wiley-Liss, Inc.     

Why are rare traits unilaterally expressed?: Trait frequency and unilateral expression for cranial nonmetric traits in humans

Based on an analysis of nonmetric trait databases from several large skeletal series in Northern Europe and South America, representing 27 bilateral traits, we report a predictable relationship between the frequency of nonmetric traits and the probability that they are expressed bilaterally. In a wider sampling of traits and populations, this study thus confirms the findings of an earlier study by Ossenberg ([1981] Am. J. Phys. Anthropol. 54:471-479), which reported the same relationship for two mandibular traits. This trend was previously explained by extending the multifactorial threshold model for discontinuous traits to incorporate either separate thresholds for unilateral or bilateral expression, or by a fuzzy threshold in which the probability of bilateral expression increases away from the median threshold value. We show that the trend is produced under the standard multifactorial threshold model for discontinuous traits simply if the within-individual or developmental instability variance remains relatively constant across the range of liability. Under this assumption, the number of individuals in which one side but not the other is pushed over the threshold for trait formation will be a larger proportion of the number of individuals expressing the trait when the trait frequency is low. As trait frequency increases, the significance of within-individual variance as a determinant of trait formation decreases relative to the genetic and among-individual environmental variance. These results have implications for interpreting nonmetric trait data as well as for understanding the prevalence of unilateral vs. bilateral expression of a wide variety of discontinuous traits, including dysmorphologies in humans.     

Quantitative genetics of cranial nonmetric traits in randombred mice: heritability and etiology

Cheverud and Buikstra (1981) demonstrated a tendency for nonmetric traits representing the number of foramina to have lower heritabilities than those representing hyperstotic or hypostotic traits in a sample of rhesus macaques. Based on this observation, Cheverud and Buikstra hypothesize that differences in the heritability of the two sets of traits may be due to differences in trait etiology. This study addresses the proposed relationship between trait heritability and etiology. Heritability values are calculated for 35 cranial nonmetric traits in a sample of 320 randombred mice using analysis of variance. The results are minimally consistent with the etiological hypothesis, but only 4 of the 35 traits showed statistically significant heritability values. These results are discussed with reference to the assumption that nonmetric traits have a strong genetic component. It is concluded that the developmental pathways that genetic variation traverses before being expressed in the form of nonmetric traits must be understood before variation in nonmetric traits can be used to its fullest potential.     

A critical review of the mean measure of divergence and Mahalanobis distances using artificial data and new approaches to the estimation of biodistances employing nonmetric traits

This article reviews the two most common distance measures employed for the calculation of biodistances based on nonmetric traits, the mean measure of divergence (MMD) and the tetrachoric Mahalanobis D² distance (TMD). In addition, two new approaches for the estimation of biodistances from nonmetric traits are proposed and assessed. The first (OMD) is based on the direct application of the Mahalanobis distance to ordinally recorded data before their transformation to binary dichotomies. The second (RMD) approximates the covariances of the Mahalanobis distance by the Pearson correlation coefficients calculated in the binary dataset. The application of all four methods to artificial datasets demonstrates that they overall provide a satisfactory estimation of the biodistance among samples especially when the number of statistically non significant distances is very limited. However, the best performance is observed by the OMD, whereas special attention should be paid to the TMD since its values might come out of an ill‐conditioned system. The influence of the number of traits, the effect of missing values, as well as the validity of the test statistics used to assess biodistance significance are also examined and discussed. Am J Phys Anthropol 157:284–294, 2015. © 2015 Wiley Periodicals, Inc.     

Cranial variation in prehistoric human skeletal remains from the Marianas

Nonmetric cranial variation and facial flatness of the Pacific and circum-Pacific populations are investigated. The peoples of the Marianas, eastern Polynesia and Hawaii form a cluster and show affinities in terms of nonmetric cranial variation with the Southeast and East Asians rather than with the Jomon-Ainu, a view which is widely supported by others. Facial flatness analysis also indicates that Polynesians have different patterns of facial prominence as compared with the Jomon-Ainu. These results increase the difficulty of accepting the Jomon-Pacific cluster proposed by Brace and his coworkers. Although genetic and nonmetric cranial variation reveal relatively close relationships, the Mariana skeletons are markedly different in facial flatness and limb bone morphology from those of Polynesians. Am J Phys Anthropol 104:399–410, 1997. © 1997 Wiley-Liss, Inc.     

Maximum-likelihood variance components analysis of heritabilities of cranial nonmetric traits

Nonmetric traits of the cranium are often used to support hypotheses of the history and divergence of human populations. These studies rely on the assumption that nonmetric traits are heritable, yet few skeletal series exist with associated pedigree information that allow for the calculation of additive genetic variance, or heritability. In addition, traits for which heritabilities have been published represent dichotomous present/absent forms instead of the range of expression that can be observed for many nonmetric characters. In the present study I use a maximum-likelihood variance components analysis to calculate univariate narrow-sense heritability estimates on the skeletal series from Hallstatt, Austria, for 9 sutural bones, 27 multilevel traits, and dichotomized present/absent forms for 19 of these multilevel characters. Most of the trait heritabilities do not differ significantly from a model of h2 = 0, and they have large standard errors. In a heuristic comparison of multilevel versus dichotomous trait forms, most of the nonmetric characters showed no differences in heritability between the two methods used for parsing the phenotypic variation, although where differences were noted, the presence-absence version had higher heritabilities. These results have implications not only for the use of particular nonmetric traits in population studies but also for the practice of character dichotomization in data collection.     

Asymmetry of the os pubis: Implications for the Suchey‐Brooks method

Studies of skeletal development frequently document populational incidences of bilateral asymmetry. Degenerative morphological skeletal changes, attributed to age related and irregular ossification, may also progress asymmetrically, either as the result of asymmetric biomechanical factors expressed over the lifespan, asymmetric expression of physiological processes, or progressive magnification of asymmetry acquired previously during development. This study illustrates the effects of bilateral asymmetry on age at death estimates obtained from human skeletal remains. The Suchey‐Brooks method, which uses the pubic symphyseal face for age estimation (Katz and Suchey, Am J Phys Anthropol 69 1986 427–435), was selected for the study based on its widespread use. Asymmetry in the Suchey‐Brooks symphyseal age phases was found in over 60% of a sample composed of 20th century White male individuals from 18 to 86 years of age (N = 130). However, results suggest that the presence of asymmetry does not compromise the accuracy of the Suchey‐Brooks method if the morphologically older symphyseal face of an asymmetric individual is used to estimate age at death. In addition, weak directional asymmetry and a correlation between age and asymmetry were found. This suggests that a comparison of asymmetry in this area with that in other skeletal areas, where the factors originating and influencing asymmetry are better understood, may be useful in better understanding the biological processes which underlie the age markers used in the Suchey‐Brooks method. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Comparability of multiple data types from the bering strait region: Cranial and dental metrics and nonmetrics,mtDNA, and Y‐chromosome DNA

Different data types have previously been shown to have the same microevolutionary patterns in worldwide data sets. However, peopling of the New World studies have shown a difference in migration paths and timings using multiple types of data, spurring research to understand why this is the case. This study was designed to test the degree of similarity in evolutionary patterns by using cranial and dental metric and nonmetric data, along with Y‐chromosome DNA and mtDNA. The populations used included Inuits from Alaska, Canada, Siberia, Greenland, and the Aleutian Islands. For comparability, the populations used for the cranial and molecular data were from similar geographic regions or had a shared population history. Distance, R and kinship matrices were generated for use in running Mantel tests, PROTEST analyses, and Procrustes analyses. A clear patterning was seen, with the craniometric data being most highly correlated to the mtDNA data and the cranial nonmetric data being most highly correlated with the Y‐chromosome data, while the phenotypic data were also linked. This patterning is suggestive of a possible male or female inheritance, or the correlated data types are affected by the same or similar evolutionary forces. The results of this study indicate cranial traits have some degree of heritability. Moreover, combining data types leads to a richer knowledge of biological affinity. This understanding is important for bioarchaeological contexts, in particular, peopling of the New World studies where focusing on reconciling the results from comparing multiple data types is necessary to move forward. Am J Phys Anthropol 154:334–348, 2014. © 2014 Wiley Periodicals, Inc.     

The quantitative genetics of fluctuating asymmetry: a comparison of two models

The genetic basis of fluctuating asymmetry (FA), a measure of random deviations from perfect bilateral symmetry, has been the subject of much recent work. In this paper we compare two perspectives on the quantitative genetic analysis of FA and directional asymmetry (DA). We call these two approaches the character-state model and the environmental responsiveness model. In the former approach, the right and left sides are viewed as separate traits whose genetic coupling is manifested by the genetic correlation. This model leads to the relationship, h2(DA) = h2[(1-rA)/(1-rp)), where h2 is the heritability of each component trait (assumed to be the same), rA and rp are the genetic and phenotypic correlations between traits, respectively. Simulation shows that, under this model, the heritability of FA is considerably less than that of DA, except when heritabilities are very close to zero. The environmental responsiveness model permits genetic variance in FA even when the genetic correlation between traits is + 1. Simulation shows that under this model the heritability of FA can be uncoupled from that of DA. The additive and nonadditive components of the component (right and left) traits, their DA and FA values are estimated using a diallel cross of seven inbred lines of the sand cricket, Gryllus firmus. Four leg measurements were made and both the individual DA and FA values and the compound measures DASUM and CFA estimated. The heritabilities of the compound measures are slightly larger than the individual estimates. Dominance variance is observed in the individual traits but predicted to be an even smaller component of the phenotypic variance than the additive genetic variance. The estimated values confirm this, although a previous study has demonstrated that dominance variance is present. Because the heritabilities of FA are generally larger than those of DA, which never exceed 0.02, the environmental responsiveness model is more consistent with the data than the character-state model. A review of other data suggests that both sources of variation might be found in some species.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. I. Single trait heritabilities

The use of skeletal nonmetric traits in studies of biological relationships often involves the assumption that variation in these traits is genetic. Studies of nonmetric traits in human groups and in inbred strains of mice and rabbits have indicated a genetic component to nonmetric trait variation. Skeletons of animals with known matrilineage membership were obtained from the Cayo Santiago skeletal collection in order to obtain a direct estimate of the heritabilities of several nonmetric traits in the free-ranging population of rhesus macaques on Cayo Santiago. Falconer's (1965) method was used to calculate heritability. Heritability estimates range from zero to one, and half of them are greater than 0.5. This indicates that there is a considerable amount of genetic variation for these traits among the Cayo macaques. There is a significant tendency for traits scoring the number of foramina to have lower heritabilities than those scoring hyperstotic or hypostotic traits.     

Brief communication: Type II tooth cusp occurrence asymmetry in a human monozygotic twin pair

A Type II tooth cusp occurrence asymmetry proposed for human twins in 1974 but not observed until recently was described in a female monozygotic twin pair. Am J Phys Anthropol 105:93–95, 1998. © 1998 Wiley-Liss, Inc.     

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.     

Twenty-five thousand-year-old triple burial from Dolní Věstonice: An ice-age family?

In 1986 a palaeolithic triple burial was discovered near Dolní Věstonice (Czech Republic). The occurrence of anatomic variants in all three skeletons gave rise to speculations that the buried individuals may have been closely related. To test this hypothesis the skeletons were submitted to a systematic kinship analysis based on odontologic and other non-metric traits. Statistical tests showed that the coincident occurrence of several rare traits in the individuals is highly unlikely to occur at random. This and further data included in the analysis therefore suggest that the three individuals buried together were genetically related and actually belonged to one family. Am J Phys Anthropol 102:123–131. © 1997 Wiley-Liss, Inc.     

Fluctuating asymmetry and sexual selection

Fluctuating asymmetry occurs when an individual is unable to undergo identical development on both sides of a bilaterally symmetrical trait. Fluctuating asymmetry measures the sensitivity of development to a wide array of genetic and environmental stresses. We propose that fluctuating asymmetry is used in many signalling contexts for assessment of an individual's ability to cope with its environment. We hypothesize that fluctuating asymmetry is used in sexual selection, both in fighting and mate choice, and in competition for access to resources. Evidence is reviewed showing that the patterns of fluctuating asymmetry in secondary sexual characters differ from those seen in other morphological traits. Secondary sexual characters show much higher levels of fluctuating asymmetry. Also, there is often a negative relationship between fluctuating asymmetry and the absolute size of ornaments, whereas the relationship is typically U-shaped in other morphological traits. The common negative relationship between fluctuating asymmetry and ornament size suggests that many ornaments reliably reflect individual quality.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. II. Phenotypic,genetic, and environmental correlations between traits

The general lack of phenotypic correlation among skeletal nonmetric traits has been interpreted as indicating a lack of genetic correlation among these traits. Nonmetric traits scored on animals in the skeletal collection of rhesus macaques from Cayo Santiago are used to calculate phenotypic, genetic, and environmental correlations between traits. The results show that even when phenotypic correlations are low, there may be large, significant genetic correlations among these traits. The genetic correlation pattern suggests that genes which affect nonmetric trait variation act primarily at a local level in the cranium, even though there are genes with pleiotropic effects on skeletal nonmetric traits throughout the cranium. Environmental and phenotypic correlations do not show this neighborhood pattern of correlation.     

Skeletal biology of Apurguan: A precontact Chamorro site on Guam

The human skeletal remains of a minimum of 152 individuals from the precontact Latte Period (AD 1000–1521) on Guam, Mariana Islands, are described. The sample, recovered at Apurguan, in the Tamuning District, is one of the largest series of well-provenienced Chamorro skeletal remains to be analyzed in recent years. The size and systematic nature of this database are a major contribution to the human biology of the region. Paleodemographic characteristics, dental and skeletal morphology, and paleopathology are presented, along with a limited examination of sex differences in frequencies of nonmetric variation. The mortuary sample, consisting of 51 subadults and 101 adults, exhibits underrepresentation of females, highest subadult mortality between 2 and 10 years, and an adult average age-at-death of 43.5 years. Cranial and infracranial indices and nonmetric variation are consistent with the Chamorro's Southeast Asian origins. There are few statistically significant sex differences in nonmetric variation which suggests close genetic affinity. The frequency of dental pathology overall is low, reflecting a well-balanced, varied diet, and consistent with preagricultural subsistence; however statistically significant sex differences suggest the influence of differential cultural behaviors or resource access. Paleopathological observations include healed fractures (more common in males), little advanced osteoarthritis, evidence for gouty arthritis, and treponemal disease (yaws). One individual, a young adult male, was interred with 10 human bone spear points in situ. Twenty percent of the primary burials exhibit evidence of postdepositional removal of selected skeletal elements for cultural purposes such as keepsakes or raw material. Am J Phys Anthropol 104:291–313, 1997. © 1997 Wiley-Liss, Inc.     

Test of revised method of age estimation from the auricular surface of the ilium

The objective of this paper is to test a revised method of age estimation based on the morphology of the auricular surface recently proposed by Buckberry and Chamberlain ([2002] Am. J. Phys. Anthropol. 119:231-239). The study sample consists of 309 individuals of known sex, age, and race from the Terry and Huntington Collections. Auricular surfaces were scored using the revised technique to determine whether it is equally applicable to both sexes as well as blacks and whites. The auricular surfaces of the same individuals were also scored using the original method of auricular surface scoring developed by Lovejoy et al. ([1985] Am. J. Phys. Anthropol. 68:15-28) to determine whether the revised technique is comparable to the original method in terms of accuracy. Results show that the revised method is equally applicable to males and females as well as blacks and whites. The revised method is less accurate than the original method for individuals between 20-49 years of age, but more accurate for individuals between 50-69 years of age.     

The repeatability of fluctuating asymmetry: a revision and extension

Fluctuating asymmetry (FA) is an imperfect measure of the developmental stability of an individual. Estimates of the heritability of FA will underestimate the heritability of developmental stability; the extent of this bias can be estimated and corrected by estimating the repeatability of developmental stability. This note corrects an error in a previous derivation of this repeatability using the absolute value of the difference between sides as the measure of FA, and derives the repeatability of developmental stability for another common measure of FA, the variance among sides within an individual.