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 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.     

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.     

The skeletal biology of two Italian peninsular Magna Graecia necropoles, Timmari and Montescaglioso

The aim of this work is to outline a general picture of life style and conditions of a population living in Magna Graecia between the 7th and the 4th c. BC by the study of human skeletal remains found in two necropoles from the Matera province, Timmari and Montescaglioso, neighbouring Metaponto, one of the main Ionian Greek colonies. The biological reconstruction was attempted by a holistic approach which foresees the use of anthropometric, anthroposcopic, palaeodemographic, palaeopathological data, the study of skeletal and dentoalveolar indicators of environmental stress and the integration with archaeological and historical information. Interpretation of the results was also based on comparisons with coeval sites from Central-Southern Italy, from Greece and with earlier and later sites from the same region. The two samples from Matera did not show appreciable differences with the other Southern Italian coeval series when compared on the basis of metric and morphometric traits. The comparison with Greek samples was hampered by the scarcity of pertinent data. A high level of muscular activity was observed in males and females, with males clearly more mobile than females. Sexual dimorphism and limb bone lateralisation were marked. Health conditions and nutritional status were good.     

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.     

Study on long bones: variation in angular traits with sex, age, and laterality

The biological problems considered in this study are sexual dimorphism in angular osteometric traits, changes in these traits with aging and their bilateral asymmetry in the same individual. The sample comprises the right and left long bones of 200 skeletons (100 males and 100 females) of Sardinian adults (Frasetto collection) for whom the sex and age at death are known. As concerns the sexual dimorphism, some angular traits of upper limb (joint axis angle of the ulna in particular) and the lower limb (condylo-diaphyseal angle of the femur and retroversion angle of the tibia) are significantly different between males and females. Torsion angles of long bones (especially the femur) were found to change with age. In sides comparison revealed a clear right-left asymmetry in our sample. The asymmetry, sexual dimorphism and changes with age are discussed in relation to varying stresses of different living activities. Our data indicate that the angular traits of long bones could be very informative in biomechanical interpretations of human activities based on skeletal remains.     

Discontinuous traits of the skull: variations on sex, age, laterality

This paper discusses some problems relating to the study and the utilization of discontinuous traits in anthropology. The importance of sex, age and laterality in the manifestation of these traits is discussed on the basis of investigations on a numerous skeletal sample of Siena (Tuscany, Italy) and on data from literature. It is pointed out that sides and sexes should be considered separately. The age of adult subjects seems to be insignificant as the incidence of these traits is concerned.     

Methodological considerations on the study of sexual dimorphism in past human populations

The degree of sexual dimorphism in human populations is influenced by stress, social role and by labour division. However, studies on ethnographic populations provided contradictory results. Unfortunately, most of these studies were based on stature only, which, as we could observe in a survey on pre-protohistoric circum-Medirerranean samples, is a poor indicator of functionally related dimorphism. A number of skeletal measurements were examined: skull, stature, transverse trunk diameters, long bones length, circumference and section, in order to assess their usefulness as indicators of functionally related dimorphism. The best indicators were represented by section and circumference of the long bones of the limbs, followed by cross-shoulder breadth (biclavicular length), stature and limb bone length, facial measurements, cranial measurements and sacral breadth. From the methodological point of view, it was found that:

1. It is better to calculate the index of dimorphism for each trait or set of traits within each sample. Then a weighted average of all the available samples is taken. The index derived from pooling a number of samples does not make biological sense. In pooled samples the distinction of between versus within sample differences is obscured;
2. It is better to combine an index which is based on the difference between averages and one which takes variability into account, because variability can also be an aspect of sexual dimorphism;
3. It is better to apply some allometric correction to the measurements used. For instance, the log transformation produces clearer trends of differential dimorphism among the various traits.

     

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.     

Selective predation of the pike Esox lucius: comparison of lateral plates and some metric features of the three-spined stickleback Gasterosteus aculeatus

Selective predation of Esox lucius on Gasterosteus aculeatus in a natural assemblage of fish of a stream in Central Poland is described and analysed. The number of lateral plates, the standard length and four size-dependent traits were significantly smaller for sticklebacks from stomach samples than for those from population samples. Thc remaining three metric traits were not significantly different among the samples. The regressions of five metric features (out of seven) on the standard length were significantly different between the sample of caten sticklebacks and that from the free-living population. This supports the supposition that the selectivity of pikc predation with respect to metric traits is not attributable to selection on body size of the prey. Thus, it is reasonable to claim that the sample of caten sticklebacks is selectively taken from the natural population. It justifies the selective predation assumption of some hypotheses of the causes of the pattern of geographic variation in the three-spined stickleback.     

Enthesopathy formation in the humerus: Data from known age‐at‐death and known occupation skeletal collections

Enthesopathies, in the guise of musculoskeletal skeletal stress markers (MSM), have been widely used to reconstruct activity levels in human skeletal populations. In general, studies have focused on their presence in the upper limb, which is used in the majority of daily activities. The aim of this study was to use some of the attachment sites on the humerus to explore the relationship between enthesopathy formation, activity, and the ageing process. The skeletal sample used in this study comprised male adult skeletons with known age‐at‐death and known occupations from the late‐19th and early‐20th century cemeteries in Portugal. The enthesopathies were recorded as either present or absent. Statistical analysis using Fishers exact tests and logistic regression was undertaken to determine whether associations could be found between specific activities or socioeconomic status (manual or nonmanual workers), and age and enthesopathy presence. Left and right sides were analyzed separately. Fisher's exact tests were used to determine the relationship between activity and enthesopathy, and they demonstrated no association between activity and enthesopathies (P > 0.01). The results of the logistic regression established that age was the single most significant factor in enthesopathy formation (P > 0.05). This study found that, in these samples, age‐at‐death, and therefore age‐related degeneration rather than degeneration caused by activities, was the primary cause of enthesopathy formation. Considering the difficulties of reliably ageing adult human skeletal remains, this is a major issue for studies of activity using enthesopathies. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

The reliability of osteometric techniques for the sex determination of burned human skeletal remains

The influence of heat-induced shrinkage on the osteometric sexual dimorphism of human skeletons is still poorly known. In order to investigate this issue, a sample composed of 84 Portuguese individuals cremated at a modern crematorium was examined using standard measurements from the femur, the talus and the calcaneus. In addition, sex determination of the sample was attempted by using osteometric standards developed from the Coimbra collection of identified skeletons. This was carried out to assess the extent of the effect of heat-induced shrinkage on the correct classification of known-sex skeletons while using standards developed on unburned skeletons.Results demonstrated that sexual dimorphism was still observable in the sample of calcined bones despite shrinkage. However, the application of conventional osteometric standards was unsuccessful. As expected, shrinkage caused most females to be correctly classified according to sex, but the sex allocation of males was very poor for all standard measurements.The results were obtained on a small sample but suggest that univariate metric techniques specifically developed for calcined bones may be valuable for sex determination. This would bring new methodological possibilities for biological anthropology and would enlarge the set of techniques regarding sex determination of burned skeletal remains.     

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.     

Activity-induced skeletal markers in historical anthropological material

A new research way of palaeopathology has recently opened: it concerns the study of the impact of the human activities on the skeleton and the reconstitution of these activities. Being interested essentially in the activity-induced skeletal markers, we carried out this study on 354 human skeletal remains. The Hungarian series represents a larger sample (263 skeletons from the 10th century; collections of the Department of Anthropology, József Attila University, Szeged, Hungary). We considered also as comparison series a collection of 91 skeletons dating from the Late Antiquity (4th century A.D.) from Provence (France), stored in the collections of the Centre Archéologique du Var (Toulon, France). The present work is based on the results of a general palaeopathological analysis with a particular attention to the possible activity-linked microtraumatic lesions. Articular and periarticular markers of activity were detected as secondary degenerative joint diseases and mechanical enthesopathies. In spite of the high rate of non-specific cases, the alterations reveal several possible activities, historically and archaeologically well documented as riding on horse-back in the Hungarian series. The sexual inegality in the distribution of the macrotraumatic lesions (male predominance of the fractures and sprains in the Hungarian series) reveals also some information about the activities of the ancient Hungarians.     

新疆哈密黑沟梁墓地出土人骨的创伤、病理及异常形态研究

本文是对出土于新疆哈密黑沟梁墓地古代人骨的病理、创伤及异常形态的综合研究。通过检查,笔者发现齿科疾病、关节疾病、创伤等病理现象在该人群中都有一定频率的出现。在对这些现象进行客观描述的基础上, 本文对这些病理现象出现的原因以及反映出该批人群的生活状况做出推论。文中同时对骨骼形态与功能的关系做了尝试性分析和讨论。     

The influence of artificial cranial deformation on discontinuous morphological traits

Following the elucidation by geneticists of the nature of minor skeletal variants in the mouse, anthropologists have stressed the potential of these traits for tracing the affinities and movements of extinct human populations. Earlier Sullivan observed that discrete traits could be particularly valuable where artificial cranial deformation limits the use of craniometry. Twenty-eight minor variants were studied in bifronto-occipitally deformed and undeformed skulls of a sample of 78 from a single Hopewell mound. The pattern of frequency differences between deformed and undeformed with respect to traits at the back of the vault and in the frontal region, interpreted in developmental terms, reveals a hypostotic effect in these regions in the deformed skull; while, in contrast, traits of the lateral vault, facial skeleton and cranial base point to a general hyperostotic effect in these regions. Each of three emissaria tends to be more constant in the deformed. That minor cranial variants manifest a plastic response to this type of environmentally-imposed stress is consistent with the nature of such variants as elucidated by genetics research in mice. The findings suggest that deformed crania should be excluded from population studies in which genetic divergence between groups is estimated in terms of cranial trait frequencies.     

Sex determination using the femur in an ancient Japanese population

Determination of sex from the femur measurements has been attempted in several populations and various studies have demonstrated the importance of population specific standards in the metric assessment of sex. The present study attempts to establish metric standards for sex determination by using femur measurements for ancient Japanese populations. Osteometric data were obtained from 151 adult skeletal remains from Jomon period, Yoshigo human skeletal collection. Eight femur measurements were taken and the data were analyzed by discriminant analysis using SPSS version 10.0. For the univariate discriminant function derived, precision of sex determination was 93% with the condyle breadth. Prediction values showed that sex differentiation could be done by femur measurements with reliability between 66.9 and 100%, with values for males higher than for females. It is suggested that discriminant formulas developed by femur measurements in this study, can be used for sex determination accurately on fragmentary skeletal remains in ancient Japanese populations.     

Dimensions of health and social structure in the early intermediate period cemetery at Villa El Salvador, Peru

This paper examines relationships between the social structure of a community and the health of its members, based on analysis of human skeletal remains (N = 64) from Villa El Salvador XII (100 BC-AD 100), a prehistoric cemetery located in the lower Lurín Valley, Peru. The ambiguity of social status as conventionally inferred from archaeological context is among the principal complicating factors in such an inquiry. We use multidimensional scaling of skeletal markers to identify the presence of patterned health-based heterogeneity in our sample, without making a priori assumptions about underlying social structure at Villa El Salvador. This procedure situates every skeleton relative to all others in the sample on the basis of multiple health markers, eliciting health groups. Once recognized, the relevance of those groups to social structure can be evaluated by comparison with a broad range of presumptive archaeological status indicators. We test the hypothesis that the distribution of stress indicators in human skeletons covaries with archaeological indicators of social differentiation. Based on multivariate analysis of skeletal indicators, we conclude that the cemetery at Villa El Salvador was utilized by two social groups with different geographic affinities: one of local coastal origin, and the other probably from the upper Lurín Valley or adjacent higher altitudes. These groups differ in skeletal characteristics related to childhood health, probably reflecting systematic contrasts in the growth environments of the studied individuals. This same division is independently supported by the distribution of cranial deformation, a possible marker of ethnicity. We also document some inequality in the distribution of labor among male individuals, as reflected by the relative advancement of degenerative joint disease, and congruent with differences in the number and quality of associated funerary offerings.