The Skeletal Phenotype of Negritos from the Andaman Islands and Philippines Relative to Global Variation among Hunter-Gatherers

The negrito hypothesis suggests that populations of small-bodied foragers in South and Southeast Asia who share common phenotypic characteristics may also share a common, ancient origin. The key defining characteristics of the negrito phenotype, small body size, dark skin, and tightly curled hair, have been interpreted as linking these populations to sub-Saharan Africans. The underlying assumption of this interpretation is that the observed phenotypic similarities likely reflect shared ancestry rather than phenotypic convergence. Current genetic evidence is inconclusive, as it both demonstrates that negrito populations have genetic affinities with neighboring populations but also rare and ancient variation that suggests considerable isolation. This study investigates the skeletal phenotype of Andaman Islanders and Aeta foragers from the Philippines in the context of the phenotypic variation among other hunter-gatherers globally, to test whether they show a common, unique physique apart from small body size. Particular emphasis is placed on the comparison of negrito phenotypes to African, Asian, and Australian hunter-gatherer diversity to investigate phenotypic similarities to other populations globally. The results demonstrate that despite sharing small adult stature, the Andaman Islanders and Aeta show variation in body dimensions. In particular, the Andaman Islanders share a pattern of narrow bi-iliac breadth and short upper limbs with the Khoisan (Later Stone Age Southern Africans), whereas the Aeta and Efé show broader bi-iliac breadths relative to lower limb lengths. Although general similarities in size and proportions remain between the Andamanese and Aeta, differences in humero-femoral indices and arm length between these groups and the Efé demonstrate that there is not a generic pygmy phenotype. Our interpretations of negrito origins and adaptation must account for this phenotypic variation

Spatial and temporal variation of body size among early Homo.

The estimation of body size among the earliest members of the genus Homo (2.4-1.5Myr [millions of years ago]) is central to interpretations of their biology. It is widely accepted that Homo ergaster possessed increased body size compared with Homo habilis and Homo rudolfensis, and that this may have been a factor involved with the dispersal of Homo out of Africa. The study of taxonomic differences in body size, however, is problematic. Postcranial remains are rarely associated with craniodental fossils, and taxonomic attributions frequently rest upon the size of skeletal elements. Previous body size estimates have been based upon well-preserved specimens with a more reliable species assessment. Since these samples are small (n < 5) and disparate in space and time, little is known about geographical and chronological variation in body size within early Homo. We investigate temporal and spatial variation in body size among fossils of early Homo using a 'taxon-free' approach, considering evidence for size variation from isolated and fragmentary postcranial remains (n = 39). To render the size of disparate fossil elements comparable, we derived new regression equations for common parameters of body size from a globally representative sample of hunter-gatherers and applied them to available postcranial measurements from the fossils. The results demonstrate chronological and spatial variation but no simple temporal or geographical trends for the evolution of body size among early Homo. Pronounced body size increases within Africa take place only after hominin populations were established at Dmanisi, suggesting that migrations into Eurasia were not contingent on larger body sizes. The primary evidence for these marked changes among early Homo is based upon material from Koobi Fora after 1.7Myr, indicating regional size variation. The significant body size differences between specimens from Koobi Fora and Olduvai support the cranial evidence for at least two co-existing morphotypes in the Early Pleistocene of eastern Africa.JTS would also like to acknowledge generous funding from the Natural Environment Research Council, UK, (Grant Number: NE/M/S/2003/00069) for supporting this research.This is the final published version. It first appeared at http://www.sciencedirect.com/science/article/pii/S0047248415000287#

Early Life Conditions and Physiological Stress following the Transition to Farming in Central/Southeast Europe: Skeletal Growth Impairment and 6000 Years of Gradual Recovery.

Early life conditions play an important role in determining adult body size. In particular, childhood malnutrition and disease can elicit growth delays and affect adult body size if severe or prolonged enough. In the earliest stages of farming, skeletal growth impairment and small adult body size are often documented relative to hunter-gatherer groups, though this pattern is regionally variable. In Central/Southeast Europe, it is unclear how early life stress, growth history, and adult body size were impacted by the introduction of agriculture and ensuing long-term demographic, social, and behavioral change. The current study assesses this impact through the reconstruction and analysis of mean stature, body mass, limb proportion indices, and sexual dimorphism among 407 skeletally mature men and women from foraging and farming populations spanning the Late Mesolithic through Early Medieval periods in Central/Southeast Europe (~7100 calBC to 850 AD). Results document significantly reduced mean stature, body mass, and crural index in Neolithic agriculturalists relative both to Late Mesolithic hunter-gatherer-fishers and to later farming populations. This indication of relative growth impairment in the Neolithic, particularly among women, is supported by existing evidence of high developmental stress, intensive physical activity, and variable access to animal protein in these early agricultural populations. Among subsequent agriculturalists, temporal increases in mean stature, body mass, and crural index were more pronounced among Central European women, driving declines in the magnitude of sexual dimorphism through time. Overall, results suggest that the transition to agriculture in Central/Southeast Europe was challenging for early farming populations, but was followed by gradual amelioration across thousands of years, particularly among Central European women. This sex difference may be indicative, in part, of greater temporal variation in the social status afforded to young girls, in their access to resources during growth, and/or in their health status than was experienced by men.Grant sponsorship Cambridge Commonwealth, European and International Trust (UK; AAM), Social Sciences and Humanities Research Council (Canada; AAM), European Research Council (RP; ERC Starting Grant, ERC-2010-StG263441), Natural Environment Research Council (JTS; NERC Grant Number NE/M/S/2003/00069).This is the final version of the article. It first appeared from PLOS via http://dx.doi.org/10.1371/journal.pone.014846

Population history and ecology, in addition to climate, influence human stature and body proportions.

Worldwide variation in human stature and limb proportions is widely accepted to reflect thermal adaptation, but the contribution of population history to this variation is unknown. Furthermore, stature and relative lower limb length (LLL) show substantial plastic responses to environmental stressors, e.g., nutrition, pathogen load, which covary with climate. Thus ecogeographic patterns may go beyond temperature-based selection. We analysed global variation in stature, sitting height and absolute and relative LLL using large worldwide samples of published anthropometric data from adult male (n = 571) and female (n = 268) populations in relation to temperature, humidity, and net primary productivity (NPP). Population history was modeled using spatial eigenvector mapping based on geographic distances reflecting the hypothesized pattern for the spread of modern humans out of Africa. Regression models account for ~ 50% of variation in most morphological variables. Population history explains slightly more variation in stature, sitting height and LLL than the environmental/climatic variables. After adjusting for population history, associations between (usually maximum) temperature and LLL are consistent with Allen's "rule" and may drive similar relationships with stature. NPP is a consistent negative predictor of anthropometry, which may reflect the growth-limiting effects of lower environmental resource accessibility (inversely related to NPP) and/or pathogen load

Divergence in male and female manipulative behaviors with the intensification of metallurgy in Central Europe.

Humeral morphology has been shown to reflect, in part, habitual manipulative behaviors in humans. Among Central European agricultural populations, long-term social change, increasing task specialization, and technological innovation all had the potential to impact patterns of habitual activity and upper limb asymmetry. However, systematic temporal change in the skeletal morphology of agricultural populations in this region has not been well-characterized. This study investigates diachronic patterns in humeral biomechanical properties and lengths among 174 adult Central European agriculturalists through the first ∼ 5400 years of farming in the region. Greater asymmetry in biomechanical properties was expected to accompany the introduction of metallurgy, particularly in males, while upper limb loading patterns were expected to be more similar between the Bronze and Iron Ages. Results revealed a divergence in the lateralization of upper limb biomechanical properties by sex between the Early/Middle Neolithic and Early/Middle Bronze Age. Neolithic females had significantly more variable properties than males in both humeri, while Bronze Age female properties became homogeneous and very symmetrical relative to the right-biased lateralization of contemporaneous males. The Bronze Age to Iron Age transition was associated with morphological change among females, with a significant increase in right-biased asymmetry and a concomitant reduction in sexual dimorphism. Relative to biomechanical properties, humeral length variation and asymmetry were low though some significant sexual dimorphism and temporal change was found. It was among females that the lateralization of humeral biomechanical properties, and variation within them, changed most profoundly through time. This suggests that the introduction of the ard and plow, metallurgical innovation, task specialization, and socioeconomic change through ∼ 5400 years of agriculture impacted upper limb loading in Central European women to a greater extent than men.The authors were funded by the following: Cambridge Commonwealth, European and International Trust (AAM) (http://www.cambridgetrust.org/); Social Sciences and Humanities Research Council of Canada (AAM) (http://www.sshrc-crsh.gc.ca/); and European Research Council Starting Grant ERC-2010-StG263,441 (RP) (http://erc.europa.eu/).This is the final published version. It first appeared at http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0112116

A trade-off between cognitive and physical performance, with relative preservation of brain function.

Debate surrounds the issue of how the large, metabolically expensive brains of Homo sapiens can be energetically afforded. At the evolutionary level, decreased investment in muscularity, adiposity and the digestive tract allow for a larger brain. Developmentally, high neo-natal adiposity and preferential distribution of resources to the brain provide an energetic buffer during times of environmental stress. Through an experimental design, we investigated the hypothesis of a trade-off involving brain and muscle at the acute level in humans. Mental performance was measured by a free-recall test, and physical performance by power output on an indoor rowing ergometer. Sixty-two male student rowers performed the two tests in isolation, and then again simultaneously. Paired samples t-tests revealed that both power output and mental performance reduced when tested together compared to in isolation (t(61) = 9.699, p < 0.001 and t(61) = 8.975, p < 0.001). Furthermore, the decrease in physical performance was greater than the decrease in mental performance (t(61) = -2.069, p = 0.043). This is the first investigation to demonstrate an acute level trade-off between these two functions, and provides support for the selfish brain hypothesis due to the relative preservation of cognitive function over physical power output. The underlying mechanism is unclear, and requires further work

Different environmental variables predict body and brain size evolution in Homo.

Increasing body and brain size constitutes a key macro-evolutionary pattern in the hominin lineage, yet the mechanisms behind these changes remain debated. Hypothesized drivers include environmental, demographic, social, dietary, and technological factors. Here we test the influence of environmental factors on the evolution of body and brain size in the genus Homo over the last one million years using a large fossil dataset combined with global paleoclimatic reconstructions and formalized hypotheses tested in a quantitative statistical framework. We identify temperature as a major predictor of body size variation within Homo, in accordance with Bergmann's rule. In contrast, net primary productivity of environments and long-term variability in precipitation correlate with brain size but explain low amounts of the observed variation. These associations are likely due to an indirect environmental influence on cognitive abilities and extinction probabilities. Most environmental factors that we test do not correspond with body and brain size evolution, pointing towards complex scenarios which underlie the evolution of key biological characteristics in later Homo

Human energetic stress associated with upregulation of spatial cognition

Objectives: Evolutionary life history theory has a unique potential to shed light on human adaptive capabilities. Ultra-endurance challenges are a valuable experimental model allowing the direct testing of phenotypic plasticity via physiological trade-offs in resource allocation. This enhances our understanding of how the body prioritizes different functions when energetically stressed. However, despite the central role played by the brain in both hominin evolution and metabolic budgeting, cognitive plasticity during energetic deficit remains unstudied. Materials: We considered human cognitive plasticity under conditions of energetic deficit by evaluating variability in performance in three key cognitive domains. To achieve this, cognitive performance in a sample of 48 athletes (m = 29, f = 19) was assessed before and after competing in multiday ultramarathons. Results: We demonstrate that under conditions of energetic deficit, performance in tasks of spatial working memory (which assessed ability to store location information, promoting landscape navigation and facilitating resource location and calorie acquisition) increased. In contrast, psychomotor speed (reaction time) remained unchanged and episodic memory performance (ability to recall information about specific events) decreased. Discussion: We propose that prioritization of spatial working memory performance during conditions of negative energy balance represents an adaptive response due to its role in facilitating calorie acquisition. We discuss these results with reference to a human evolutionary trajectory centred around encephalisation. Encephalisation affords great plasticity, facilitating rapid responses tailored to specific environmental conditions, and allowing humans to increase their capabilities as a phenotypically plastic species

A trade-off between cognitive and physical performance, with relative preservation of brain function

Debate surrounds the issue of how the large, metabolically expensive brains of Homo sapiens can be energetically afforded. At the evolutionary level, decreased investment in muscularity, adiposity and the digestive tract allow for a larger brain. Developmentally, high neo-natal adiposity and preferential distribution of resources to the brain provide an energetic buffer during times of environmental stress. Through an experimental design, we investigated the hypothesis of a trade-off involving brain and muscle at the acute level in humans. Mental performance was measured by a free-recall test, and physical performance by power output on an indoor rowing ergometer. Sixty-two male student rowers performed the two tests in isolation, and then again simultaneously. Paired samples t-tests revealed that both power output and mental performance reduced when tested together compared to in isolation (t(61) = 9.699, p < 0.001 and t(61) = 8.975, p < 0.001). Furthermore, the decrease in physical performance was greater than the decrease in mental performance (t(61) = -2.069, p = 0.043). This is the first investigation to demonstrate an acute level trade-off between these two functions, and provides support for the selfish brain hypothesis due to the relative preservation of cognitive function over physical power output. The underlying mechanism is unclear, and requires further work