Adult proportionality in small-bodied foragers: a test of ecogeographic expectations

If predictable, ecogeographic patterning in body size and proportions of human populations can provide valuable information regarding human biology, adaptation to local environments, migration histories, and health, now and in the past. This paper evaluates the assumption that small-bodied Later Stone Age (LSA) foragers of Southern Africa show the adult proportions that would be expected of warm-adapted populations. Comparisons are also made with small-bodied foragers from the Andaman Islands (AI). Indices including brachial, crural, limb element length to skeletal trunk height, and femoral head and bi-iliac breadth to femoral length were calculated from samples of LSA (n = 124) and AI (n = 31) adult skeletons. Samples derived from the literature include those from high (Europe), middle (North Africa), and low (Sub-Saharan Africa) latitude regions. The LSA and AI samples match some but not all expected ecogeographic patterns for their particular regions of long term habitation. For most limb length to skeletal trunk height indices the LSA and AI are most similar to the other mid-latitude sample (North Africans). However, both groups are similar to low latitude groups in their narrow bi-iliac breadths, and the AI display relatively long radii. Proportions of LSA and AI samples also differ from those of African pygmies. In regions like southern-most Africa, that do not experience climatic extremes of temperature or humidity, or where small body size exists through drift or selection, body size, and proportions may also be influenced by nonclimatic variables, such as energetic efficiency.     

Hunter-gatherer postcranial robusticity relative to patterns of mobility, climatic adaptation, and selection for tissue economy

Human skeletal robusticity is influenced by a number of factors, including habitual behavior, climate, and physique. Conflicting evidence as to the relative importance of these factors complicates our ability to interpret variation in robusticity in the past. It remains unclear how the pattern of robusticity in the skeleton relates to adaptive constraints on skeletal morphology. This study investigates variation in robusticity in claviculae, humeri, ulnae, femora, and tibiae among human foragers, relative to climate and habitual behavior. Cross-sectional geometric properties of the diaphyses are compared among hunter-gatherers from southern Africa (n = 83), the Andaman Islands (n = 32), Tierra del Fuego (n = 34), and the Great Lakes region (n = 15). The robusticity of both proximal and distal limb segments correlates negatively with climate and positively with patterns of terrestrial and marine mobility among these groups. However, the relative correspondence between robusticity and these factors varies throughout the body. In the lower limb, partial correlations between polar second moment of area (J(0.73)) and climate decrease from proximal to distal section locations, while this relationship increases from proximal to distal in the upper limb. Patterns of correlation between robusticity and mobility, either terrestrial or marine, generally increase from proximal to distal in the lower and upper limbs, respectively. This suggests that there may be a stronger relationship between observed patterns of diaphyseal hypertrophy and behavioral differences between populations in distal elements. Despite this trend, strength circularity indices at the femoral midshaft show the strongest correspondence with terrestrial mobility, particularly among males.     

Interpreting skeletal growth in the past from a functional and physiological perspective

The study of juvenile skeletal remains can yield important insights into the health, behavior, and biological relationships of past populations. However, most studies of past skeletal growth have been limited to relatively simple metrics. Considering additional skeletal parameters and taking a broader physiological perspective can provide a more complete assessment of growth patterns and environmental and genetic effects on those patterns. We review here some alternative approaches to ontogenetic studies of archaeological and paleontological skeletal material, including analyses of body size (stature and body mass) and cortical bone structure of long bone diaphyses and the mandibular corpus. Together such analyses can shed new light on both systemic and localized influences on bone growth, and the metabolic and mechanical factors underlying variation in growth. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

Robusticity and sexual dimorphism in the postcranium of modern hunter-gatherers from Australia

Throughout much of prehistory, humans practiced a hunting and gathering subsistence strategy. Elevated postcranial robusticity and sexually dimorphic mobility patterns are presumed consequences of this strategy, in which males are attributed greater robusticity and mobility than females. Much of the basis for these trends originates from populations where skeletal correlates of activity patterns are known (e.g., cross-sectional geometric properties of long bones), but in which activity patterns are inferred using evidence such as archaeological records (e.g., Pleistocene Europe). Australian hunter-gatherers provide an opportunity to critically assess these ideas since ethnographic documentation of their activity patterns is available. We address the following questions: do skeletal indicators of Australian hunter-gatherers express elevated postcranial robusticity and sexually dimorphic mobility relative to populations from similar latitudes, and do ethnographic accounts support these findings. Using computed tomography, cross-sectional images were obtained from 149 skeletal elements including humeri, radii, ulnae, femora, and tibiae. Cross-sectional geometric properties were calculated from image data and standardized for body size. Australian hunter-gatherers often have reduced robusticity at femoral and humeral midshafts relative to forager (Khoi-San), agricultural/industrialized (Zulu), and industrialized (African American) groups. Australian hunter-gatherers display more sexual dimorphism in upper limb robusticity than lower limb robusticity. Attributing specific behavioral causes to upper limb sexual dimorphism is premature, although ethnographic accounts support sex-specific differences in tool use. Virtually absent sexual dimorphism in lower limb robusticity is consistent with ethnographic accounts of equivalently high mobility among females and males. Thus, elevated postcranial robusticity and sexually dimorphic mobility do not always characterize hunter-gatherers.     

Levels of Intraspecific Variation Within the Catarrhine Skeleton

The amount of phenotypic variation between conspecifics is largely a result of the opposing forces of constraint and plasticity. Because selection is the product of competition between individuals of the same species, understanding the interactions between these forces is vital to understanding evolution. We investigated levels of intraspecific variation in the catarrhine skeleton using a morphometric analysis of 245 crania and 189 appendicular postcranial skeletons. We identified regions of interest from the literature and made comparisons of intraspecific variation between the appendicular postcranium and cranium, the forelimb and hind limb, the diaphyses and epiphyses of long bones, and the diaphyses of the proximal and distal segments of the limbs. We confirmed that variation is significantly higher in the appendicular postcranium than in the cranium, in the forelimb compared to the hind limb, and in the diaphyses compared to the epiphyses. Further, we found that this pattern was repeated in 12 species spanning the infraorder, suggesting a characteristic of catarrhines in general. The relatively low level of variation in the cranium suggests that constraint is more widespread in this region compared to the appendicular postcranium, which is more plastic, especially in the diaphyses of the forelimb. In contrast to previous studies, we found the diaphyses of the distal segment to be more variable than the diaphyses of the proximal segment. The results from this study, which show regional differences in intraspecific variation, will aid the interpretation of evolutionary and plastic influences on morphological variation and inform decisions about which skeletal regions are suitable for answering specific evolutionary questions.     

Articular and diaphyseal remodeling of the proximal femur with changes in body mass in adults.

Proximal femoral dimensions were measured from radiographs of 80 living subjects whose current body weight and body weight at initial skeletal maturity (18 years) could be ascertained. Results generally support the hypothesis that articular size does not change in response to changes in mechanical loading (body weight) in adults, while diaphyseal cross-sectional size does. This can be explained by considering the different bone remodeling constraints characteristic of largely trabecular bone regions (articulations) and largely compact cortical bone regions (diaphyses). The femoral neck shows a pattern apparently intermediate between the two, consistent with its structure. When the additional statistical "noise" created by an essentially static femoral head size is accounted for, the present study supports other studies that have demonstrated rather marked positive allometry in femoral articular and shaft cross-sectional dimensions to body mass among adult humans. Body weight prediction equations developed from these data give reasonable results for modern U.S. samples, with average percent prediction errors of about 10%-16% for individual weights and about 2% for sample mean weights using the shaft dimension equations. When predicting body weight from femoral head size in earlier human samples, a downward correction factor of about 10% is suggested to account for the increased adiposity of very recent U.S. adults.     

Sequential changes in weight of the skeleton and in length of long limb bones of Macaca mulatta.

In a collection of 274 monkeys (Macaca mulatta) the relative weight of the dry, fat-free skeleton, expressed as a proportion of total body weight, increases significantly throughout the gestational period to approximately 6% with only random variation after birth. The weight of the fetal skeleton increases exponentially with age. In the postnatal period the skeletal weight increases asymptotically to adulthood, which is considered to be 6.5 years of age. Equations for estimating skeletal weight are presented. Of four subdivisions of the skeleton, the skul contributes the greatest proportion of total skeletal weight in the fetal stage with the proportion decreasing to adulthood. The contributions of the other subdivisions, postcranial axial, superior limb, and inferior limb, and inferior limb, are nearly equal in the fetal stage, with that of only the inferior limb increasing to adulthood, when it makes up the greatest proportion of total skeletal weight. Until the last third of the gestational period, the humerus is longer than the femur and the radius longer than the tibia. Thereafter, the inferior limbs grow at a faster rate than the superior limbs, resulting in an intermembral index of approximately 95% by birth and less than 90% by adulthood.     

Brief communication: The effects of disuse on the mechanical properties of bone: What unloading tells us about the adaptive nature of skeletal tissue

The intricate link between load environment and skeletal health is exemplified by the severe osteopenia that accompanies prolonged periods of immobilization, frequently referred to as disuse osteoporosis. Investigating the effects disuse has on the structural properties of bone provides a unique opportunity to better understand how mechanical loads influence the adaptation and maintenance of skeletal tissue. Here, we report results from an examination of multiple indicators of bone metabolism (e.g., mean osteon density, mean osteon size, bone mass, and bone area distribution) within the major long bones of individuals with distinct activity level differences. Results are based on a sample comprising two subjects that suffered from long‐term quadriplegia and 28 individuals of comparable age that had full limb mobility. Although limited in sample size, our findings suggest bones associated with long‐term disuse have lower osteon densities and larger osteon areas compared to individuals of normal mobility, reflecting dramatically lower remodeling rates potentially related to reduced strain levels. Moreover, immobilized skeletal elements demonstrate a reduced percentage of cortical area present resulting from endosteal resorption. Differences between mobility groups in the percentage of cortical area present and bone distribution of all skeletal elements, suggests bone modeling activity is negligible in the unloaded adult skeleton. Additional histomorphometric comparisons reveal potential intraskeletal differences in bone turnover rates suggesting remodeling rates are highest within the humeri and femora. Addition of more immobilized individuals in the future will allow for quantitative statistical analyses and greater consideration of human variation within and between individuals. Am J Phys Anthropol 2012. © 2012 Wiley Periodicals, Inc.     

Intraskeletal variability in bone mass

For methodological or other reasons, a variety of skeletal elements are analyzed and subsequently used as a basis for describing general bone loss and mass. However, bone loss and mass may not be uniform within and among skeletal elements of the same individual because of biomechanical factors. We test the hypothesis that a homogeneity in bone mass exists among skeletal elements of the same individual. Measures indicative of bone mass were calculated from the midshafts of six skeletal elements from the same individuals (N = 41). The extent of intraskeletal variability in bone mass (relative cortical area) was then examined for the entire sample, according to age, sex, and pathological status. The results of the analysis showed that all measures reflect a heterogeneity in bone mass (P 相似文献   

Skeletal growth in early and late Neolithic foragers from the Cis‐Baikal region of Eastern Siberia

Skeletal growth is explored between Early Neolithic (EN) (8000 to 6800 BP) and Late Neolithic (LN) (6000 to 5200 BP) foragers from the Cis‐Baikal region of Eastern Siberia. Previous studies suggest that increased systemic stress and smaller adult body size characterize the EN compared to LN. On this basis, greater evidence for stunting and wasting is expected in the EN compared to LN. Skeletal growth parameters assessed here include femoral and tibial lengths, estimated stature and body mass, femoral midshaft cortical thickness, total bone thickness, and medullary width. Forward selection was used to fit polynomial lines to each skeletal growth parameter relative to dental age in the pooled samples, and standardized residuals were compared between groups using t tests. Standardized residuals of body mass and femoral length were significantly lower in the EN compared to LN sample, particularly from late infancy through early adolescence. However, no significant differences in the standardized residuals for cortical thickness, medullary width, total bone thickness, tibial length, or stature were found between the groups. Age ranges for stunting in femoral length and wasting in body mass are consistent with environmental perturbations experienced at the cessation of breast feeding and general resource insecurity in the EN compared to LN sample. Differences in relative femoral but not tibial length may be associated with age‐specific variation in growth‐acceleration for the distal and proximal limb segments. Similarity in cortical bone growth between the two samples may reflect the combined influences of systemic and mechanical factors on this parameter. Am J Phys Anthropol 153:377–386, 2014. © 2013 Wiley Periodicals, Inc.     

Mechanical determinants of bone form: insights from skeletal remains

Analysis of skeletal remains from humans living in the past forms an important complement to observational and experimental studies of living humans and animal models. Including earlier humans in such analyses increases the range of variation in both behavior and body size and shape that are represented, and can provide insights into the adaptive potential of the modern human skeleton. I review here a variety of studies of archaeological and paleontological remains that have investigated differences in skeletal structure from a mechanical perspective, focusing in particular on diaphyseal strength of the limb bones. Several conclusions can be drawn from these studies: 1) there has been a decline in overall skeletal strength relative to body size over the course of human evolution that has become progressively steeper in recent millennia, probably due to increased sedentism and technological advancement; 2) differences in pelvic structure and hip mechanical loadings affect femoral shape; 3) activity patterns affect overall strength and shape of both the lower and upper limb bones; and 4) responsiveness to changes in mechanical loading varies between skeletal features (e.g., articulations versus diaphyses) and by age.     

Postcranial morphology and locomotion of the Eocene raoellid Indohyus (Artiodactyla: Mammalia)

Raoellids are small, raccoon-sized Eocene artiodactyls, closely related to archaic cetaceans (archaeocetes) that have poor representation of postcranial elements in the fossil record. Little is known of the aquatic and terrestrial locomotor affinities of raoellids due to the paucity of their fossil record, leaving a critical gap in our understanding of the earliest portion of the artiodactyl marine invasion. To address this gap, a comparative morphological analysis of the postcranial elements was undertaken based on newly recovered elements of the raoellid Indohyus, archaeocetes and extant artiodactyls. Greater than 200 postcranial elements of Indohyus were described, and some limb element cross-sections were visualised via paleohistological thin sections and CT scans. Results show that during terrestrial locomotion, Indohyus probably had a digitigrade posture and mediolaterally stabilised limbs that functioned mostly in flexion and extension within the parasagittal plane. Quantification of midshaft cross-sectional area for some elements of Indohyus showed an osteosclerotic cortex, a skeletal characteristic associated with aquatic behaviours. Indohyus may represent a critical intermediate in the evolution of the cetacean terrestrial-to-aquatic body plan, as it bears gracile postcranial element proportions similar to a terrestrial artiodactyl but also an incipient form of osteosclerosis compared to pakicetid archaeocetes.     

Protection of obstetric dimensions in a small-bodied human sample

In human females, the bony pelvis must find a balance between being small (narrow) for efficient bipedal locomotion, and being large to accommodate a relatively large newborn. It has been shown that within a given population, taller/larger-bodied women have larger pelvic canals. This study investigates whether in a population where small body size is the norm, pelvic geometry (size and shape), on average, shows accommodation to protect the obstetric canal. Osteometric data were collected from the pelves, femora, and clavicles (body size indicators) of adult skeletons representing a range of adult body size. Samples include Holocene Later Stone Age (LSA) foragers from southern Africa (n = 28 females, 31 males), Portuguese from the Coimbra-identified skeletal collection (CISC) (n = 40 females, 40 males) and European-Americans from the Hamann-Todd osteological collection (H-T) (n = 40 females, 40 males). Patterns of sexual dimorphism are similar in the samples. Univariate and multivariate analyses of raw and Mosimann shape-variables indicate that compared to the CISC and H-T females, the LSA females have relatively large midplane and outlet canal planes (particularly posterior and A-P lengths). The LSA males also follow this pattern, although with absolutely smaller pelves in multivariate space. The CISC females, who have equally small stature, but larger body mass, do not show the same type of pelvic canal size and shape accommodation. The results suggest that adaptive allometric modeling in at least some small-bodied populations protects the obstetric canal. These findings support the use of population-specific attributes in the clinical evaluation of obstetric risk.     

Diaphysator: An online application for the exhaustive cartography and user-friendly statistical analysis of long bone diaphyses

The cross-sectional geometry (CSG) of long bone diaphyses is used in bioanthropology to evaluate their resistance to biomechanical constraints and to infer life-history-related patterns such as mobility, activity specialization or intensity, sexual dimorphism, body mass and proportions. First limited by technical analytical constraints to the analysis of one or two cross sections per bone, it has evolved into the analysis of cross sections of the full length of the diaphyseal part of long bones. More recently, researchers have developed analytical tools to map the cortical thickness of entire diaphyses to evaluate locomotor signatures. However, none of these analytical tools are easy to use for scientists who are not familiar with computer programming, and some statistical procedures–such as mapping the correlation coefficients of the diaphyseal thickness with various parameters have yet to be made available. Therefore, we developed an automated and open-source application that renders those analyses (both CSG and cortical thickness) in a semiautomated and user friendly manner. This application, called “Diaphysator”, is associated with another free software (“Extractor”, presented in Dupej et al. (2017). American Journal of Physical Anthropology, 164, 868–876). Diaphysator can be used as an online application ( https://diaphysator.shinyapps.io/maps ) or as a package for R statistical software. Along with the mean maps of cortical thickness and mean CSG parameter graphs, the users can evaluate the correlations and partial correlations of both CSG parameters at every cross section along the diaphyseal length, and cortical thickness data points of the entire diaphysis, with any factor such as age, sex, stature, and body mass.     

Influence of loading condition and anatomical location on human cortical bone linear micro-cracks

Human cortical bone fracture toughness depends on the anatomical locations under quasi-static loading. Recent results also showed that under fall-like loading, cortical bone fracture toughness is similar at different anatomical locations in the same donor. While cortical bone toughening mechanisms are known to be dependent on the tissue architecture under quasi-static loading, the fracture mechanisms during a fall are less studied. In the current study, the structural parameters of eight paired femoral diaphyses, femoral necks and radial diaphyses were mechanically tested under quasi-static and fall-like loading conditions (female donors, 70 ± 14 y.o., [50–91 y.o.]). Synchrotron radiation micro-CT imaging was used to quantify the amount of micro-cracks formed during loading. The volume fraction of these micro-cracks was significantly higher within the specimens loaded under a quasi-static condition than under a loading representative of a fall. Under fall-like loading, there was no difference in crack volume fraction between the different paired anatomical locations. This result shows that the micro-cracking toughening mechanism depends both on the anatomical location and on the loading condition.     

Pulmonary pneumaticity in the postcranial skeleton of extant aves: a case study examining Anseriformes

Anseriform birds were surveyed to examine how the degree of postcranial pneumaticity varies in a behaviorally and size-diverse clade of living birds. This study attempts to extricate the relative effects of phylogeny, body size, and behavioral specializations (e.g., diving, soaring) that have been postulated to influence the extent of postcranial skeletal pneumaticity. One hundred anseriform species were examined as the focal study group. Methods included latex injection of the pulmonary apparatus followed by gross dissection or direct examination of osteological specimens. The Pneumaticity Index (PI) is introduced as a means of quantifying and comparing postcranial pneumaticity in a number of species simultaneously. Phylogenetically independent contrasts (PICs) were used to examine the relationship between body size and the degree of postcranial pneumaticity throughout the clade. There is a high degree of similarity (i.e., clade-specificity) within most anseriform subgroups. As a whole, Anseriformes demonstrate no significant relationship between relative pneumaticity and body size, as indicated by regression analysis of body mass on PI. It is apparent, however, that many clades of diving ducks do exhibit lower PIs than their nondiving relatives. By exclusion of diving taxa from analyses, a significant positive slope is observed and the hypothesis of relatively higher pneumaticity in larger-bodied birds is only weakly supported. However, low correlations indicate that factors other than body size account for much of the variation observed in relative pneumaticity. Pneumaticity profiles were mapped onto existing phylogenetic hypotheses. A reduction in the degree of postcranial pneumaticity occurred independently in at least three anseriform subclades specialized for diving. Finally, enigmatic pneumatic features located in distal forelimb elements of screamers (Anhimidae) result from invasion of bone by a network of subcutaneous air sac diverticula spreading distally along the wings.     

Immobilization and bone structure in humans

Long-term immobilization is known to result in substantial bone loss. The present review examined the existing evidence for deterioration of bone structure during long-term disuse in humans. Paralysis due to spinal cord injury, long-term exposure to microgravity in space or tightly restricted mobility during bed rest provide reasonable models to assess the influence of immobilization on bone structure. Expectedly, the duration of immobilisation was the major determinant of bone loss, but irrespective of whether the skeletal unloading was due to irrecoverable paralysis, long-term spaceflight or bed rest, the mean pattern of structural deterioration of bone, mainly manifest as substantial cortical thinning and trabecular bone loss, was quite similar. However, skeletal responses to disuse can be highly variable between individuals. Apparently the relative decline in individual’s bone loading in relation to loading prior to immobilization accounts for inter-individual variation.     

Body size, body shape, and long bone strength in modern humans

To identify behaviorally significant differences in bone structure it is first necessary to control for the effects of body size and body shape. Here the scaling of cross-sectional geometric properties of long bone diaphyses with different "size" measures (bone length, body mass, and the product of bone length and body mass) are compared in two modern human populations with very different body proportions: Pecos Pueblo Amerindians and East Africans. All five major long bones (excluding the fibula) were examined. Mechanical predictions are that cortical area (axial strength) should scale with body mass, while section modulus (bending/torsional strength) should scale with the product of body mass and moment arm length. These predictions are borne out for section moduli, when moment arm length is taken to be proportional to bone length, except in the proximal femoral diaphysis, where moment arm length is proportional to mediolateral body breadth (as would be expected given the predominance of M-L bending loads in this region). Mechanical scaling of long bone bending/torsional strength is similar in the upper and lower limbs despite the fact that the upper limb is not weight-bearing. Results for cortical area are more variable, possibly due to a less direct dependence on mechanical factors. Use of unadjusted bone length alone as a "size" measure produces misleading results when body shape varies significantly, as is the case between many modern and fossil hominid samples. In such cases a correction factor for body shape should be incorporated into any "size" standardization.     

F-81 skeleton from Wadi Mataha, Jordan, and its bearing on human variability in the Epipaleolithic of the Levant

The discovery of a Middle Epipaleolithic adult skeleton (F-81) at the site of Wadi Mataha in southern Jordan provides new insights into human variability in the Epipaleolithic of the Levant. This paper analyzes the skeletal morphology of Wadi Mataha F-81 in the context of other Epipaleolithic remains from Jordan and Israel to assess the current evidence for morphological variability throughout this period. The F-81 skeleton shares morphological features with earlier Epipaleolithic skeletons from Ohalo and Nahal Ein Gev, and later Natufian populations. Despite the morphological similarities, F-81 extends the range of known variability prior to the Natufian with its unusually small stature and unique combination of morphological characteristics. High levels of cranial and postcranial robusticity suggest that the F-81 individual was physically active and terrestrially mobile. Pronounced bilateral asymmetry in the upper limb suggests significant lateralization of habitual activity. In the context of Epipaleolithic remains, the F-81 skeleton provides preliminary evidence for greater morphological variability, terrestrial mobility, and lateralized habitual behavior prior to the Natufian, and skeletal gracilization between the Middle and Late Epipaleolithic in the Levant.     

Skeletal indicators of subsistence patterns and activity régime in the Mesolithic sample from Grotta dell’Uzzo (Trapani,Sicily): a case study

The skeletal sample from Grotta dell’Uzzo, Sicily (2 adult females and 5 adult males) was compared to a number of more representative population samples from Western Europe and the Mediterranean Basin. The majority of these were from Italian pre- and protohistoric sites. The research protocol analyzed skeletal indicators of labour activity and sexual division of labour (body size and proportions, sexual dimorphism, limb lateralization, bone robustness, the development of muscular attachments, accessory articular facets, signs of muscular hyperfunction). Sexual dimorphism and limb lateralization showed some regular patterns of possible general significance in all the samples examined here. A general pattern of gracilization and de-specialization of physical activity is observed in the Mesolithic as compared to the Upper Palaeolithic samples. The main features of the Mesolithic samples are: a reduction of body size and bone robustness, a lower degree of sexual dimorphism and limb bone asymmetry, a reduction of the mechanical stress on the lower limbs indicated by less pronounced muscular attachments and reduced talar flattening. This trend is reversed towards the Neolithic period. The main features of these variations are discussed in relation to economic and environmental changes. The Uzzo sample fits well into the general picture of the Western European Mesolithic, although showing some intermediate features between the Mesolithic and the Neolithic samples. This paper is dedicated to the memory of the late Roland Menk, who made significant contribution to our understanding of the Mesolithic transition.