Mobility and subsistence economy: a diachronic comparison between two groups settled in the same geographical area (Liguria, Italy)

The purpose of this article is to investigate temporal shifts in skeletal robusticity to infer behavioral changes in two populations (Neolithic, NEOL and Medieval, MED) settled in the same geographic area but involved in different subsistence economies (pastoralism and coastal resources exploitation). This comparison allows us to test the hypothesis that occupational stress and mobility in the same environment produce predictable changes in the robusticity of both upper and lower limbs. Results show a lower degree of humeral robusticity and a similar degree of humeral asymmetry in the two sexes in the MED population. These results are consistent with the relatively less stressful subsistence economy documented in the MED population relative to that of the NEOL. Lower limb results suggest that femoral robusticity does not correlate directly with the level of logistical mobility, but is instead due to the summation of several diverse factors that place biomechanical loads on the hindlimb, particularly unevenness of the terrain. However, female femoral gracility seems to indicate that below a certain "threshold" of mobility, i.e., movement over the natural terrain, terrain conformation is no longer the main contributing factor to femoral robusticity. The femoral shape index I(x)/I(y) declines through time, particularly in males. This agrees with the expected mobility of the samples based on archaeological and historical data, providing further evidence on the reliability of this index in inferring terrestrial mobility.     

Mouse hallucal metatarsal cross‐sectional geometry in a simulated fine branch niche

Mice raised in experimental habitats containing an artificial network of narrow “arboreal” supports frequently use hallucal grasps during locomotion. Therefore, mice in these experiments can be used to model a rudimentary form of arboreal locomotion in an animal without other morphological specializations for using a fine branch niche. This model would prove useful to better understand the origins of arboreal behaviors in mammals like primates. In this study, we examined if locomotion on these substrates influences the mid‐diaphyseal cross‐sectional geometry of mouse metatarsals. Thirty CD‐1/ICR mice were raised in either arboreal (composed of elevated narrow branches of varying orientation) or terrestrial (flat ramps and walkways that are stratified) habitats from weaning (21 days) to adulthood (≥4 months). After experiments, the hallucal metatarsal (Mt1) and third metatarsal (Mt3) for each individual were isolated and micro‐computed tomography (micro‐CT) scans were obtained to calculate mid‐shaft cross‐sectional area and polar section modulus. Arboreal mice had Mt1s that were significantly more robust. Mt3 cross sections were not significantly different between groups. The arboreal group also exhibited a significantly greater Mt1/Mt3 ratio for both robusticity measures. We conclude that the hallucal metatarsal exhibits significant phenotypic plasticity in response to arboreal treatment due to habitual locomotion that uses a rudimentary hallucal grasp. Our results support the hypothesis that early adaptive stages of fine branch arboreality should be accompanied by a slightly more robust hallux associated with the biomechanical demands of this niche. J. Morphol. 276:759–765, 2015. © 2015 Wiley Periodicals, Inc.     

Human manipulative behavior in the Central European Late Eneolithic and Early Bronze Age: humeral bilateral asymmetry

It is assumed that the transition from the Late Eneolithic to the Early Bronze Age in Central Europe was associated with substantial changes in subsistence and the perception of gender differences. However, the archeological record itself does not entirely support this model. Alternatively, this transition may be interpreted as a continuous process. We used asymmetry in external dimensions, and asymmetry in size and distribution of cortical tissue of humeri to elucidate the nature of this transition with respect to differences in manipulative behavior. The total sample of 67 individuals representing five archaeological cultures was used. The results indicate that the pattern of asymmetry of the humeral external measurements and the cross-sectional parameters taken at 35% of humeral biomechanical length remain stable during the Late Eneolithic and Early Bronze Age. However, females of both periods show fluctuating asymmetry for all of the cross-sectional parameters, but directional asymmetry for biomechanical length. Males are nonsignificantly shifted from the line of equivalence for biomechanical length, but exhibit directional asymmetry for the cortical area and polar moment of area. Only distal articular breadth yields fluctuating asymmetry for both females and males in both periods. Thus, the transition from the Late Eneolithic to the Early Bronze Age can be seen as a continuous process that probably affected only a limited part of human activities. We interpret the differences between females and males of both periods as evidence of gender-specific activities; males might have been associated with extra-domestic agricultural labor that resulted in asymmetrical manipulative loading and females with domestic labor with symmetrical manipulative loading in both periods.     

Biomechanical approach to the reconstruction of activity patterns in Neolithic Western Liguria, Italy

This paper investigates the changes in upper and lower limb robusticity and activity patterns that accompanied the transition to a Neolithic subsistence in western Liguria (Italy). Diaphyseal robusticity measures were obtained from cross-sectional geometric properties of the humerus and femur in a sample of 16 individuals (eight males and eight females) dated to about 6,000-5,500 BP. Comparisons with European Late Upper Paleolithics (LUP) indicate increased humeral robusticity in Neolithic Ligurian (NEOL) males, but not in females, with a significant reduction in right-left differences in both sexes. Sexual dimorphism in robusticity increases in upper and lower limb bones. Regarding the femur, while all female indicators of bending strength decrease steadily through time, values for NEOL males approach those of LUP. This suggests high, and unexpected, levels of mechanical stress for NEOL males, probably reflecting the effects of the mountainous terrain on lower limb remodeling. Comparisons between NEOL males and a small sample of LUP hunter-gatherers from the same area support this interpretation. In conclusion, cross-sectional geometry data indicate that the transition to Neolithic economies in western Liguria did not reduce functional requirements in males, and suggest a marked sexual division of labor involving a more symmetrical use of the upper limb, and different male-female levels of locomotory stress. When articulated with archaeological, faunal, paleopathological, and ethnographic evidence, these results support the hypothesis of repetitive, bimanual use of axes tied to pastoral activities in males, and of more sedentary tasks linked to agriculture in females.     

Modeling and remodeling responses to normal loading in the human lower limb

Limb bones are designed to be strong enough to support the body and yet be energetically conservative during locomotion. Bones of the distal segment, which are relatively costly to move, are often more slender than bones of the proximal segments, even though they must sustain proportionally greater loads. As a result, they are expected to experience a higher incidence of microdamage. With this constraint in mind, Lieberman and Crompton (1998 Principles of Animal Design, Cambridge: Cambridge University Press, p. 78-86) proposed that bones response to strain varies along the proximo-distal axis of the limb. In order to avoid fatigue fractures due to the accumulation of microdamage, the distal segment, in comparison to the proximal segment, will have an increase in remodeling events to replace damaged bone. In this paper, we test the hypothesis of Lieberman and Crompton (1998) with respect to the human lower limb. With a sample of adult individuals, we compare tibiae and femora for mid-diaphyseal cross-sectional geometry and Haversian remodeling differences. Our results indicate that the human limb is not designed like that of quadrupedal cursorial animals. The tibia is not less resistant in bending and torsion, and does not remodel more than the femur. Our findings fail to support the hypothesis of Lieberman and Crompton (1998) and suggest, instead, that the human lower limb is not designed like a cursorial animal limb. In addition, our results support previous observations that remodeling is not uniform within the cross section of a bone, probably a reflection of different loading histories within the different regions of the cross section.     

Declining tibial curvature parallels ∼6150 years of decreasing mobility in central european agriculturalists

Long bones respond to mechanical loading through functional adaptation in a suite of morphological characteristics that together ensure structural competence to in vivo loading. As such, adult bone structure is often used to make inferences about past behavior from archaeological remains. However, such biomechanical approaches often investigate change in just one aspect of morphology, typically cross‐sectional morphology or trabecular structure. The relationship between longitudinal bone curvature and mobility patterns is less well understood, particularly in the tibia, and it is unknown how tibial curvature and diaphyseal cross‐sectional geometry interact to meet the structural requirements of loading. This study examines tibial curvature and its relationship with diaphyseal cross‐sectional geometry (CSG) and body size in preindustrial Central Europeans spanning ～6150 years following the introduction of agriculture in the region. Anteroposterior centroid displacement from the proximo‐distal longitudinal axis was quantified at nine diaphyseal section locations (collectively representative of diaphyseal curvature) in 216 tibial three‐dimensional laser scans. Results documented significant and corresponding temporal declines in midshaft centroid displacement and CSG properties. Significant correlations were found between mid‐diaphyseal centroid displacement and all mobility‐related CSG properties, while the relationship weakened toward the diaphyseal ends. No significant relationship was found between centroid displacement and body size variables with the exception of the most distal section location. Results support a relationship between tibial curvature and cross‐sectional geometry among prehistoric Central European agricultural populations, and suggest that changes in mechanical loading may have influenced a suite of morphological features related to bone adaptation in the lower limb. Am J Phys Anthropol 157:260–275, 2015. © 2015 Wiley Periodicals, Inc.     

Cross sectional geometry of the forelimb skeleton and flight mode in pelecaniform birds

Avian wing elements have been shown to experience both dorsoventral bending and torsional loads during flapping flight. However, not all birds use continuous flapping as a primary flight strategy. The pelecaniforms exhibit extraordinary diversity in flight mode, utilizing flapping, flap‐gliding, and soaring. Here we (1) characterize the cross‐sectional geometry of the three main wing bone (humerus, ulna, carpometacarpus), (2) use elements of beam theory to estimate resistance to loading, and (3) examine patterns of variation in hypothesized loading resistance relative to flight and diving mode in 16 species of pelecaniform birds. Patterns emerge that are common to all species, as well as some characteristics that are flight‐ and diving‐mode specific. In all birds examined, the distal most wing segment (carpometacarpus) is the most elliptical (relatively high I_max/I_min) at mid‐shaft, suggesting a shape optimized to resist bending loads in a dorsoventral direction. As primary flight feathers attach at an oblique angle relative to the long axis of the carpometacarpus, they are likely responsible for inducing bending of this element during flight. Moreover, among flight modes examined the flapping group (cormorants) exhibits more elliptical humeri and carpometacarpi than other flight modes, perhaps pertaining to the higher frequency of bending loads in these elements. The soaring birds (pelicans and gannets) exhibit wing elements with near‐circular cross‐sections and higher polar moments of area than in the flap and flap‐gliding birds, suggesting shapes optimized to offer increased resistance to torsional loads. This analysis of cross‐sectional geometry has enhanced our interpretation of how the wing elements are being loaded and ultimately how they are being used during normal activities. J. Morphol., 2011. © 2011 Wiley‐Liss,Inc.     

Technical note: The effect of midshaft location on the error ranges of femoral and tibial cross‐sectional parameters

In comparing long‐bone cross‐sectional geometric properties between individuals, percentages of bone length are often used to identify equivalent locations along the diaphysis. In fragmentary specimens where bone lengths cannot be measured, however, these locations must be estimated more indirectly. In this study, we examine the effect of inaccurately located femoral and tibial midshafts on estimation of geometric properties. The error ranges were compared on 30 femora and tibiae from the Eneolithic and Bronze Age. Cross‐sections were obtained at each 1% interval from 60 to 40% of length using CT scans. Five percent of deviation from midshaft properties was used as the maximum acceptable error. Reliability was expressed by mean percentage differences, standard deviation of percentage differences, mean percentage absolute differences, limits of agreement, and mean accuracy range (MAR) (range within which mean deviation from true midshaft values was less than 5%). On average, tibial cortical area and femoral second moments of area are the least sensitive to positioning error, with mean accuracy ranges wide enough for practical application in fragmentary specimens (MAR = 40–130 mm). In contrast, tibial second moments of area are the most sensitive to error in midshaft location (MAR = 14–20 mm). Individuals present significant variation in morphology and thus in error ranges for different properties. For highly damaged fossil femora and tibiae we recommend carrying out additional tests to better establish specific errors associated with uncertain length estimates. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Linking structural variability in long bone diaphyses to habitual behaviors: foragers from the southern African Later Stone Age and the Andaman Islands

The cross-sectional distribution of cortical bone in long bone diaphyses is highly responsive to mechanical loading during life, yet the relationship between systemic and localized influences on skeletal structure remains unclear. This study investigates postcranial robustness throughout the body among adults from two groups of foragers with different patterns and modes of mobility, to determine whether there is evidence for upper vs. lower body localization of skeletal robustness. The samples used for this comparison are from the southern African Later Stone Age (LSA; n = 65, male = 33, female = 28) dating from ca. 10,000 to 2,000 B.P., and 19th century indigenous Andaman Islanders (AI; n = 36, male = 17, female = 16). The LSA were highly mobile foragers who did not exploit offshore marine resources. In contrast, the AI had tightly constrained terrestrial, but significant marine, mobility. Geometric properties of cortical bone distribution in the diaphyses of the clavicle, humerus, femur, tibia, and first metatarsal are compared between the samples, providing a representation of skeletal robustness throughout the body. Multivariate ANOVA shows the AI to have significantly stronger clavicles and humeri, while the LSA femora, tibiae, and first metatarsals are stronger than those of the AI. These patterns, in which upper and lower limbs show biomechanical properties that are consistent with habitual behaviors, suggest localized osteogenic response. Although postcranial robustness appears to be correlated with overall limb function, the results suggest that more proximal elements within the limb may be more responsive to mechanical loading.     

The influence of body proportions on femoral and tibial midshaft shape in hunter‐gatherers

Variation in femoral and tibial diaphyseal shape is used as an indicator of adaptation to patterns of terrestrial mobility. Recent experimentation has implied that lower limb diaphyseal shape may be primarily influenced by lower limb length, and less so by mobility patterns. If valid, this would, at most, render previous interpretations of mobility patterns based on analyses of diaphyseal shape questionable, and, at least, require additional standardization that considers the influence of limb length. Although the consequences could be profound, this implication has yet to be directly tested. Additionally, the influence of body breadth on tibial shape (and to a lesser extent femoral shape) remains uncertain. Tibial and femoral cross‐sectional midshaft shape measurements, taken from nine Pleistocene and Holocene skeletal populations, were compared against lower limb length, limb segment length, and bi‐iliac breadth. Generally, limb length and limb segment length do not significantly influence femoral or tibial midshaft shape. After controlling for body mass greater bi‐iliac breadth is associated with a relative mediolateral strengthening of the femoral midshaft, while the influence of a wider body shape (BIB/length) is associated with a relative M‐L strengthening of the tibia and femur of males, and the tibia of females. We conclude that; (1) mechanical interpretations of lower limb diaphyseal shape are most parsimonious due to the lack of evidence for a consistent relationship between segment length and shape; however, (2) further work is required to investigate the influence of bi‐iliac breadth on both femoral and tibial midshaft shape. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Technical note: A stereological analysis of the cross‐sectional variability of the femoral osteon population

Unbiased selection of regions of interest (ROIs) and unbiased definition of histological structures are needed to improve the repeatability of microscopic methods for age at death determination and to reduce operator subjectivity. We present results obtained by selecting ROIs according to stereological principles on a sample of 28 femoral cross sections of Caucasoid males aged 20–89 years. A regular grid was overlaid on the cross section, and the ROIs were selected as close as possible to the periosteum in the anterior, lateral, and medial regions. The areas consisting of all intact secondary osteons plus fragments were outlined and osteon population density, percent osteon population, area, and perimeter were calculated using stereological methods and software. Overall, the analyses of intra‐ and inter‐section variability showed no significant difference between the ROIs, i.e., the location within the cross section of the ROIs does not affect the outcome of the analyses. The individual variability was found to be higher in adults aged 30–55 years than in other age ranges. ranges. Am J Phys Anthropol 2010. © 2010 Wiley‐Liss, Inc.     

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.