A Late Pleistocene human humerus from Rusinga Island, Lake Victoria, Kenya.

In 2010, a hominin right humerus fragment (KNM-RU 58330) was surface collected in a small gully at Nyamita North in the Late Pleistocene Wasiriya Beds of Rusinga Island, Kenya. A combination of stratigraphic and geochronological evidence suggests the specimen is likely between ∼49 and 36 ka in age. The associated fauna is diverse and dominated by semiarid grassland taxa. The small sample of associated Middle Stone Age artifacts includes Levallois flakes, cores, and retouched points. The 139 mm humeral fragment preserves the shaft from distal to the lesser tubercle to 14 mm below the distal end of the weakly projecting deltoid tuberosity. Key morphological features include a narrow and weakly marked pectoralis major insertion and a distinctive medial bend in the diaphysis at the deltoid insertion. This bend is unusual among recent human humeri but occurs in a few Late Pleistocene humeri. The dimensions of the distal end of the fragment predict a length of 317.9 ± 16.4 mm based on recent samples of African ancestry. A novel method of predicting humeral length from the distance between the middle of the pectoralis major and the bottom of the deltoid insertion predicts a length of 317.3 mm ± 17.6 mm. Cross-sectional geometry at the midshaft shows a relatively high percentage of cortical bone and a moderate degree of flattening of the shaft. The Nyamita humerus is anatomically modern in its morphology and adds to the small sample of hominins from the Late Pleistocene associated with Middle Stone Age artifacts known from East Africa. It may sample a population closely related to the people of the out-of-Africa migration.

An examination of the cross-sectional geometrical properties of the long bone diaphyses of Holocene foragers from Roonka, South Australia.

OBJECTIVES: This study examines long bone diaphyseal rigidity and shape of hunter-gatherers at Roonka to make inferences about subsistence strategies and mobility of inhabitants of semi-arid southeastern Australia. Roonka is a cemetery site adjacent to the Lower Murray River, which contains over 200 individuals buried throughout the Holocene. Archaeological evidence indicates that populations living near this river corridor employed mobile, risk averse foraging strategies. METHODS: This prediction of lifestyle was tested by comparing the cross-sectional geometric properties of the humerus, radius, ulna, femur, tibia, and fibula of individuals from Roonka to samples of varying subsistence strategies. Bilateral asymmetry of the upper limb bones was also examined. RESULTS: Roonka males and females have moderately high lower limb diaphyseal rigidity and shape. In the upper limb, females have low rigidity and bilateral asymmetry while males have moderately high rigidity and bilateral asymmetry. This pattern is similar to other foraging groups from Australia and southern Africa that have behaviorally adapted to arid and semi-arid environments. DISCUSSION: Lower limb results suggest that populations in the Lower Murray River Valley had relatively elevated foraging mobility. Upper limb rigidity and bilateral asymmetry indicate a sexual division of labor at Roonka. Females resemble other samples that had mixed subsistence strategies that involved hunting, gathering, and processing tasks. Males display a pattern similar to groups that preferentially hunted large game, but that supplemented this source with smaller game and riverine resources.

Are human hands and feet affected by climate? A test of Allen's rule.

OBJECTIVES: In recent years, several studies have shown that populations from cold, high-latitude regions tend to have relatively shorter limbs than populations from tropical regions, with most of the difference due to the relative length of the zeugopods (i.e., radius, ulna, tibia, fibula). This pattern has been explained either as the consequence of long-term climatic selection or of phenotypic plasticity, with temperature having a direct effect on bone growth during development. The aims of this study were to test whether this pattern of intra-limb proportions extended to the bones of the hands and feet, and to determine whether the pattern remained significant after taking into account the effects of neutral evolutionary processes related to population history. MATERIALS AND METHODS: Measurements of the limb bones, including the first metatarsal and metacarpal, were collected for 393 individuals from 10 globally distributed human populations. The relationship between intra-limb indices and minimum temperature was tested using generalized least squares regression, correcting for spatial autocorrelation. RESULTS: The results confirmed previous observations of a temperature-related gradient in intra-limb proportions, even accounting for population history. This pattern extends to the hands, with populations from cold regions displaying a relatively shorter and stockier first metacarpal; however, the first metatarsal appears to be wider but not shorter in cold-adapted populations. DISCUSSION: The results suggest that climatic adaptation played a role in shaping variation in limb proportions between human populations. The different patterns shown by the hands and feet might be due to the presence of evolutionary constraints on the foot to maintain efficient bipedal locomotion.

Developmental plasticity in fat patterning of Ache children in response to variation in interbirth intervals: a preliminary test of the roles of external environment and maternal reproductive strategies.

A firm link between small size at birth and later more centralized fat patterning has been established in previous research. Relationships between shortened interbirth intervals and small size at birth suggest that maternal energetic prioritization may be an important, but unexplored determinant of offspring fat patterning. Potential adaptive advantages to centralized fat storage (Baker et al., 2008: In: Trevathan W, McKenna J, Smith EO, editors. Evolutionary Medicine and Health: New Perspectives. New York: Oxford) suggest that relationships with interbirth intervals may reflect adaptive responses to variation in patterns of maternal reproductive effort. Kuzawa (2005: Am J Hum Biol 17:5-21; 2008: In: Trevathan W, McKenna J, Smith EO, editors. Evolutionary Medicine and Health: New Perspectives. New York: Oxford) has argued that maternal mediation of the energetic quality of the environment is a necessary component of developmental plasticity models invoking predictive adaptive responses (Gluckman and Hanson 2004: Trends Endocrinol Metab 15:183-187). This study tested the general hypothesis that shortened interbirth intervals would predict more centralized fat patterning in offspring. If long-term maternally mediated signals are important determinants of offspring responses, then we expected to observe a relationship between the average interbirth interval of mothers and offspring adiposity, with no relationship with the preceding interval. Such a finding would suggest that maternal, endogenous resource allocation decisions are related to offspring physiology in a manner consistent with Kuzawa's description. We observed exactly such a relationship among the Ache of Paraguay, suggesting that maternally mediated in utero signals of postnatal environments may be important determinants of later physiology. The implications of these findings are reviewed in light of life history and developmental plasticity theories and ourability to generalize the results to other populations. Recommendations for further empirical research are briefly summarized.

The aging of Wolff's "law": ontogeny and responses to mechanical loading in cortical bone.

The premise that bones grow and remodel throughout life to adapt to their mechanical environment is often called Wolff's law. Wolff's law, however, is not always true, and in fact comprises a variety of different processes that are best considered separately. Here we review the molecular and physiological mechanisms by which bone senses, transduces, and responds to mechanical loads, and the effects of aging processes on the relationship (if any) between cortical bone form and mechanical function. Experimental and comparative evidence suggests that cortical bone is primarily responsive to strain prior to sexual maturity, both in terms of the rate of new bone growth (modeling) as well as rates of turnover (Haversian remodeling). Rates of modeling and Haversian remodeling, however, vary greatly at different skeletal sites. In addition, there is no simple relationship between the orientation of loads in long bone diaphyses and their cross-sectional geometry. In combination, these data caution against assuming without testing adaptationist views about form-function relationships in order to infer adult activity patterns from skeletal features such as cross-sectional geometry, cortical bones density, and musculo-skeletal stress markers. Efforts to infer function from shape in the human skeleton should be based on biomechanical and developmental models that are experimentally tested and validated.

Exercise-induced bone formation is poorly linked to local strain magnitude in the sheep tibia.

Functional interpretations of limb bone structure frequently assume that diaphyses adjust their shape by adding bone primarily across the plane in which they are habitually loaded in order to minimize loading-induced strains. Here, to test this hypothesis, we characterize the in vivo strain environment of the sheep tibial midshaft during treadmill exercise and examine whether this activity promotes bone formation disproportionately in the direction of loading in diaphyseal regions that experience the highest strains. It is shown that during treadmill exercise, sheep tibiae were bent in an anteroposterior direction, generating maximal tensile and compressive strains on the anterior and posterior shaft surfaces, respectively. Exercise led to significantly increased periosteal bone formation; however, rather than being biased toward areas of maximal strains across the anteroposterior axis, exercise-related osteogenesis occurred primarily around the medial half of the shaft circumference, in both high and low strain regions. Overall, the results of this study demonstrate that loading-induced bone growth is not closely linked to local strain magnitude in every instance. Therefore, caution is necessary when bone shaft shape is used to infer functional loading history in the absence of in vivo data on how bones are loaded and how they actually respond to loading.

Abrasive, silica phytoliths and the evolution of thick molar enamel in primates, with implications for the diet of Paranthropus boisei.

BACKGROUND: Primates--including fossil species of apes and hominins--show variation in their degree of molar enamel thickness, a trait long thought to reflect a diet of hard or tough foods. The early hominins demonstrated molar enamel thickness of moderate to extreme degrees, which suggested to most researchers that they ate hard foods obtained on or near the ground, such as nuts, seeds, tubers, and roots. We propose an alternative hypothesis--that the amount of phytoliths in foods correlates with the evolution of thick molar enamel in primates, although this effect is constrained by a species' degree of folivory. METHODOLOGY/PRINCIPAL FINDINGS: From a combination of dietary data and evidence for the levels of phytoliths in plant families in the literature, we calculated the percentage of plant foods rich in phytoliths in the diets of twelve extant primates with wide variation in their molar enamel thickness. Additional dietary data from the literature provided the percentage of each primate's diet made up of plants and of leaves. A statistical analysis of these variables showed that the amount of abrasive silica phytoliths in the diets of our sample primates correlated positively with the thickness of their molar enamel, constrained by the amount of leaves in their diet (R(2) = 0.875; p<.0006). CONCLUSIONS/SIGNIFICANCE: The need to resist abrasion from phytoliths appears to be a key selective force behind the evolution of thick molar enamel in primates. The extreme molar enamel thickness of the teeth of the East African hominin Paranthropus boisei, long thought to suggest a diet comprising predominantly hard objects, instead appears to indicate a diet with plants high in abrasive silica phytoliths.

A description of the Omo I postcranial skeleton, including newly discovered fossils.

Recent fieldwork in the Kibish Formation has expanded our knowledge of the geological, archaeological, and faunal context of the Omo I skeleton, the earliest known anatomically modern human. In the course of this fieldwork, several additional fragments of the skeleton were recovered: a middle manual phalanx, a distal manual phalanx, a right talus, a large and a small fragment of the left os coxae, a portion of the distal diaphysis of the right femur that conjoins with the distal epiphysis recovered in 1967, and a costal fragment. Some researchers have described the original postcranial fragments of Omo I as anatomically modern but have noted that a variety of aspects of the specimen's morphology depart from the usual anatomy of many recent populations. Reanalysis confirms this conclusion. Some of the unusual features in Omo I--a medially facing radial tuberosity, a laterally flaring facet on the talus for the lateral malleolus, and reduced dorsovolar curvature of the base of metacarpal I--are shared with Neandertals, some early modern humans from Skhul and Qafzeh, and some individuals from the European Gravettian, raising the possibility that Eurasian early modern humans inherited these features from an African predecessor rather than Neandertals. The fragment of the os coxae does not unambiguously diagnose Omo I's sex: the greater sciatic notch is intermediate in form, the acetabulum is large (male?), and a preauricular sulcus is present (female?). The preserved portion of the left humerus suggests that Omo I was quite tall, perhaps 178-182 cm, but the first metatarsal suggests a shorter stature of 162-173 cm. The morphology of the auricular surface of the os coxae suggests a young adult age.

Further new hominin fossils from the Kibish Formation, southwestern Ethiopia.

In addition to the new fragments of the Omo I skeleton, renewed fieldwork in the Kibish Formation along the lower reaches of the Omo River in southwestern Ethiopia has yielded new hominin finds from the Kibish Formation. The new finds include four heavily mineralized specimens: a partial left tibia and a fragment of a distal fibular diaphysis from Awoke's Hominid Site (AHS), a parietal fragment, and a portion of a juvenile occipital bone. The AHS tibia and fibula derive from Member I and are contemporaneous with Omo I and II. The other specimens derive from Chad's Hominid Site (CHS), and derive from either Member III or IV, which constrains their age between approximately 8.6 and approximately 104 ka.

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.

Optimization of bone growth and remodeling in response to loading in tapered mammalian limbs.

How bones respond dynamically to mechanical loading through changes in shape and structure is poorly understood, particularly with respect to variations between bones. Structurally, cortical bones adapt in vivo to their mechanical environments primarily by modulating two processes, modeling and Haversian remodeling. Modeling, defined here as the addition of new bone, may occur in response to mechanical stimuli by altering bone shape or size through growth. Haversian remodeling is thought to be an adaptation to repair microcracks or prevent microcrack propagation. Here, we examine whether cortical bone in sheep limbs modulates periosteal modeling and Haversian remodeling to optimize strength relative to mass in hind-limb midshafts in response to moderate levels of exercise at different growth stages. Histomorphometry was used to compare rates of periosteal growth and Haversian remodeling in exercised and sedentary treatment groups of juvenile, subadult and young adult sheep. In vivo strain data were also collected for the tibia and metatarsal midshafts of juvenile sheep. The results suggest that limb bones initially optimize responses to loading according to the varying power requirements associated with adding mass at different locations. In juveniles, exercise induces higher rates of periosteal modeling in proximal midshafts and higher rates of Haversian remodeling in distal midshafts. Consequently, distal element midshafts experience higher strains and, presumably, have lower safety factors. As animals age, periosteal modeling rates decline and Haversian remodeling rates increase, but moderate levels of mechanical loading stimulate neither process significantly.