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.     

乌鲁木齐萨恩萨依墓地出土头骨的人种学研究

本文研究了乌鲁木齐萨恩萨依墓地出土的人头骨材料, 这些材料大部分属于公元前七世纪左右的墓葬。通过观察其颅、面部特点,对其颅、面部进行测量分析, 结果显示萨恩萨依墓地出土头骨的人种总体来说属于蒙古人种类型。     

Calcaneus secundarius: description and frequency in six skeletal samples

The calcaneus secundarius (CS) is a small accessory ossicle of the anterior calcaneal facet. A sample of 1,367 calcanei including early 20th century U.S. blacks and whites, XIIth Dynasty Egyptians, Islamic Period Bahrainis, and protohistoric American Indians from two states were examined to determine the frequency of this nonmetric trait. Statistical testing of the correlation of the trait with sex could only be performed on the blacks and whites, a documented cadaver collection. Tests of chi 2 indicated that, although sex differences were insignificant, significant group (not necessarily racial or geographical) differences existed at the .05 level. There was no correlation of side dominance in the overall sample, although significant population differences existed. The trait frequencies of the black and white samples were more similar to one another than to the nonindustrial groups who, in turn, were more similar to each other than to the Terry sample. The findings suggest that a CS is of genetic or congenital origin resulting in a secondary center of ossification in the anterior calcaneal facet.     

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.     

An examination of the meaning of cranial discrete traits for human skeletal biological studies

Discrete traits are of increasing interest in comparative skeletal biological research. Characteristics justifying their use have been investigated primarily in mice, however. Using 72 discrete variants, 321 human skulls from the Terry Collection of known race, sex and age have been studied. Significant sex and age differences were detected. Inter-trait correlation was found to be at a low but significant overall level. Multivariate comparison with conventional craniometric analysis was undertaken on subdivisions of the sample, and distance based on metric and nonmetric data were concordant. It is concluded, on the basis of these findings and the discontinuous variant frequency distributions, that discrete traits in isolation are not of paramount value to skeletal genetic studies, but may be vital in comparison and conjunction with other types of data in analyzing the population genetics of extinct groups.     

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.     

The supra-acetabular fossa and groove: a skeletal marker for Northwest European mesolithic populations

A nonmetric trait, known as the supra-acetabular fossa and groove, is described for European Upper Paleolithic and mesolithic populations. This morphological feature consists of a pit posterior or superior to the anterior-inferior iliac spine and an associated groove which arcs over the roof of the acetabulum. Presence of the trait seems to be primarily related to the origin of the reflected head of rectus femoris. The trait is variable in its expression, shows an increase in definition with age, and occurs equally in males and females. Although the trait is commonly found in pre-Upper Paleolithic hominids, it is extremely rare in the Upper Paleolithic, limited to a single occurrence in the Oberkassel 2 female. In the Mesolithic, except for two innominates from Vlasac (Yugoslavia), the trait is found only in NW European populations, especially Skateholm (Sweden) and Henriksholm-Bøgebakken (Denmark) where it occurs in more than 50% of the known pelves. Given the trait’s restricted geographic distribution and high occurrence in Skateholm and Henriksholm-Bøgebakken, it is suggested the supra-acetabular fossa and groove represents a skeletal marker for NW European Mesolithic populations. As such, it provides evidence for the development of social territories in the early Holocene.     

Cranial deformation and nonmetric trait variation

Cranial deformation is known to influence many traditional craniometric variables, but its effects on nonmetric trait variation are not well characterized. In this study, we examine the effects of three types of deformation (annular, lambdoid flattening, and fronto-occipital) on nonmetric traits, using a large sample of protohistoric and prehistoric crania. Our results indicate that a few traits are increased or decreased in relative frequency by particular types of deformation, but that these effects have little impact on the calculation of biological distances between groups. © 1993 Wiley-Liss, Inc.     

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.     

The relationship between cranial metric and nonmetric traits in the rhesus macaques from Cayo Santiago

This study addresses the relationship between cranial metric variables and nonmetric traits using the skeletal sample of rhesus macaques from Cayo Santiago. Discriminant function analysis is used to study the metric differences between macaque crania grouped according to the presence or absence of nine nonmetric traits. The computation of total structure coefficients from the discriminant function analyses provides information regarding how closely each metric variable is related to the discriminant functions derived. Total structure coefficients have not been used previously in the study of the relationship between metric and nonmetric traits. The results of the analysis are interpreted using an explicit approach to cranial morphogenesis-functional cranial analysis. It is concluded that the relationship between cranial metric and nonmetric traits is explicable in terms of a common developmental pathway shared by the two types of traits. Identification of the specific etiology of nonmetric traits depends on future anatomical studies or organisms throughout the period of nonmetric trait development.     

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.     

A comparative study of metric and non-metric traits in a series of modern crania

The purpose of this study is to question the utility of non-metric traits as predictors of age, race, and sex by comparing a group of non-metric variables with a group of metric variables. Twelve metric variables and 15 nonmetric variables were scored on 317 crania from the Terry Collection at the U. S. National Museum, and later statistically evaluated with a number of multivariate analysis programs and a stepwise discriminant program. The metric variables were found to be significant sex and race discriminators, where-as the non-metric variables were non-significant. The non-metric variables were found to be better age discriminators than the metric variables, even though neither could be classified as highly significant. It was concluded that non-metric traits by themselves have very little discriminatory value and should be used instead to supplement other osteological measurements and observations.     

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.     

Genetic correlations between sides and heritability of asymmetry for nonmetric traits in rhesus macaques on Cayo Santiago

The use of nonmetric traits for estimation of biological distance is a long-standing practice in biological anthropology. Nonmetric traits can be scored using either the individual or the side of the individual as the unit of measure. If sides of the individual are genetically correlated the use of sides would produce redundant genetic information. For this reason, Korey (Am. J. Phys. Anthropol. 53:19-23, 1980) argues for the use of individuals as the unit of measure for nonmetric traits. Ossenberg (Am. J. Phys, Anthropol. 54:471-479, 1981), however, argues that bilateral occurrence of nonmetric traits indicates greater genetic liability for the trait and that therefore the sides are the more biologically correct unit of measure. Genetic correlations for 13 cranial nonmetric traits are estimated for a sample of rhesus macaque skeletons from Cayo Santiago. In addition, heritability of asymmetry is estimated for these 13 traits as a test of Ossenberg's contention that asymmetry is genetically influenced. Significant genetic correlations between sides support Korey's contention that nonmetric traits should be scored by individual. Only two asymmetry heritabilities were significantly different from zero, providing no significant support for Ossenberg's contention that asymmetry is genetically determined. Our results support the theory that asymmetry represents a measure of the ability of an organism to buffer stresses. Therefore, a measure of the heritability of asymmetry is a measure of the heritability of the ability to buffer stresses. This ability does not appear to be heritable in this sample.     

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.     

Articular surface defects in the third metatarsal and third cuneiform: Nonosseous tarsal coalition

Frequencies of articular surface defects on the third metatarsal and third cuneiform, seen as pits of varying sizes on the plantar one third of the tarsometatarsal articular face, were investigated in skeletal populations from North America and Japan, as well as in gibbon, orangutan, chimpanzee, and gorilla skeletons. The apes did not exhibit the defects, although the number of observed specimens of each type was small. The newly presented human frequencies corresponded well with those from other published sources. The defects appeared both unilaterally and bilaterally, with no apparent sex or side biases. Statistical tests between the various populations found that, in general, geographically close populations had more similar frequencies of the defect. Possible etiologies for the defect were investigated, including biomechanical influences, degenerative arthritis, infection, trauma, and a developmental condition known as tarsal coalition, which proved to be the best explanation. Tarsal coalition results from the failure of a joint space to form properly during fetal growth. It can occur between any two adjacent bones of the foot. Several clinically important coalitions, whose presence interferes with normal walking, are known. However, coalition between the third metatarsal and third cuneiform has not been reported in the clinical literature, suggesting that the defect causes little or no foot dysfunction. Tarsal coalition is thought to have a strong genetic component, suggesting that the pit defect may be useful as a skeletal nonmetric trait, as others have stated. Am J Phys Anthropol 109:53–65, 1999. © 1999 Wiley-Liss, Inc.     

The effects of human disturbance on the species composition,species diversity and functional diversity of a Miombo woodland in northern Malawi

Anthropogenic disturbances have serious impacts on ecosystems across the world. Understanding the effects of disturbance on woodlands, especially in regions where local people depend on these natural resources, is essential for sustainable natural resource management and biodiversity conservation. In this study, we evaluated the effects of anthropogenic disturbance, specifically selective logging of Brachystegia floribunda, on woodlands by comparing species composition, species diversity and functional diversity of woody plants between disturbed and undisturbed woodlands. We combined species data and functional trait data for leaves, fruits and other traits related to resource and disturbance responses to calculate functional indices (functional richness, evenness and divergence) and community‐weighted means of each trait. Shifts in taxonomic species composition were analysed using nonmetric multi‐dimensional scaling. Species composition differed significantly between disturbed and undisturbed woodlands. Tree density was greater in disturbed woodlands, whereas evenness, functional evenness and functional divergence were greater in undisturbed woodlands. In terms of forest cover, selective logging of B. floribunda appeared to have little impact on Miombo woodlands, but some shifts in functional traits, such as the shift from a deciduous to evergreen phenology, may increase the vulnerability of these ecosystems to environmental change, especially drought.     

Sexing skulls using discriminant function analysis of visually assessed traits

The accuracy of sex determinations based on visual assessments of the mental eminence, orbital margin, glabellar area, nuchal area, and mastoid process was tested on a series of 304 skulls of known age and sex from people of European American, African American, and English ancestry as well as on an ancient Native American sample of 156 individuals whose sex could be reliably determined based on pelvic morphology. Ordinal scores of these sexually dimorphic traits were used to compute sex determination discriminant functions. Linear, kth-nearest-neighbor, logistic, and quadratic discriminant analysis models were evaluated based on their capacity to minimize both misclassifications and sex biases in classification errors. Logistic regression discriminant analysis produced the best results: a logistic model containing all five cranial trait scores correctly classified 88% of the modern skulls with a negligible sex bias of 0.1%. Adding age at death, birth year, and population affinity to the model did not appreciably improve its performance. For the ancient Native American sample, the best logistic regression model assigned the correct pelvic sex to 78% of the individuals with a sex bias of only 0.2%. Similar cranial trait frequency distributions were found in same-sex comparisons of the modern African American, European American, and English samples. The sexual dimorphism of these modern people contrasts markedly with that of the ancient Native Americans. Because of such population differences, discriminant functions like those presented in this paper should be used with caution on populations other than those for which they were developed.     

Logistic analysis of the effects of shovel trait on Carabelli's trait in a mongoloid population

Mongoloid populations differ from Caucasoids by having a high prevalence of shovel trait and a low prevalence of Carabelli's trait. This study was conducted to investigate the effects of the shovel trait on Carabelli's trait in a Mongoloid population. The research design sought a population that resides in an isolated area and exhibits low admixture with neighboring populations. The Mongoloid group selected for study was the Bunun tribe of aborigines who inhabit an alpine area in Taiwan. The effects of sex and age on Carabelli's trait were controlled in this investigation, as was the association between tooth size and Carabelli's trait. Results show that males were more likely to have Carabelli's trait expressed on teeth than females. The buccolingual diameter of Carabelli's trait teeth was larger than that of teeth without the trait. After adjusting for sex, age, and tooth size, the existence of the shovel trait increased the likelihood of having Carabelli's trait by a factor of three, an effect that is significant. © 1996 Wiley-Liss, Inc.     

The use of nonmetric variation in estimating human population admixture: a test case with Brazilian blacks, whites, and mulattos

Measurements in populations which serve as valid indicators of biological relationship should be proportional to genetic distance. In order to test the utility of discrete cranial traits for estimating genetic distances among populations, estimates of admixture are obtained for gene frequency data and nonmetric cranial data in São Paulo mulattos (M). The gene frequency data serve as a control that the three populations are related as stated: estimates of admixture are obtained by using São Paulo whites (W) and blacks (B) as parental populations and by estimating the parameter of admixture, m, in the model p_M = (1 ? m) p_W + mp_B (Elston, 1971) where the p's are either gene frequencies or nonmetric trait frequencies. A test of goodness of fit of the model provides a means of ascertaining whether or not the data fit this linear model. While the gene frequency data indicate distances among the three populations which are highly compatible with the linear model of admixture, the nonmetric data show significant deviations from the model. This implies that the frequencies of the nonmetric traits in the populations used in this analysis are not a linear function of genetic distance. This discourages the use of nonmetric traits in making quantitative conclusions about genetic relationships. It also suggests the need for investigation of the use of other skeletal characters for estimating genetic distance, as well as approaches for such investigations through the study of hybrid individuals.