The relative importance of ancient demography and climate in determining worldwide patterns of human within-population phenotypic diversity is still open to debate. Several morphometric traits have been argued to be under selection by climatic factors, but it is unclear whether climate affects the global decline in morphological diversity with increasing geographical distance from sub-Saharan Africa. Using a large database of male and female skull measurements, we apply an explicit framework to quantify the relative role of climate and distance from Africa. We show that distance from sub-Saharan Africa is the sole determinant of human within-population phenotypic diversity, while climate plays no role. By selecting the most informative set of traits, it was possible to explain over half of the worldwide variation in phenotypic diversity. These results mirror those previously obtained for genetic markers and show that 'bones and molecules' are in perfect agreement for humans.

The extent to which past climate change has dictated the pattern and timing of the out-of-Africa expansion by anatomically modern humans is currently unclear [Stewart JR, Stringer CB (2012) Science 335:1317-1321. In particular, the incompleteness of the fossil record makes it difficult to quantify the effect of climate. Here, we take a different approach to this problem; rather than relying on the appearance of fossils or archaeological evidence to determine arrival times in different parts of the world, we use patterns of genetic variation in modern human populations to determine the plausibility of past demographic parameters. We develop a spatially explicit model of the expansion of anatomically modern humans and use climate reconstructions over the past 120 ky based on the Hadley Centre global climate model HadCM3 to quantify the possible effects of climate on human demography. The combinations of demographic parameters compatible with the current genetic makeup of worldwide populations indicate a clear effect of climate on past population densities. Our estimates of this effect, based on population genetics, capture the observed relationship between current climate and population density in modern huntergatherers worldwide, providing supporting evidence for the realism of our approach. Furthermore, although we did not use any archaeological and anthropological data to inform the model, the arrival times in different continents predicted by our model are also broadly consistent with the fossil and archaeological records. Our framework provides the most accurate spatiotemporal reconstruction of human demographic history available at present and will allow for a greater integration of genetic and archaeological evidence.human dispersals | colonization | population bottlenecks | net primary productivity | most recent common ancestor

Recent applications of population genetic models to human craniodental traits have revealed a strong neutral component to patterns of global variation. However, little work has been undertaken to determine whether neutral processes might also be influencing the postcranium, perhaps due to substantial evidence for selection and plastic environmental responses in these regions. Recent work has provided evidence for neutral effects in the pelvis, but has been limited in regard to shape data (small numbers of linear measurements) and restricted only to males. Here, we use geometric morphometric methods to examine population variation in the human os coxae (pelvic bone) in both males and females. Neutrality is examined via apportionment of variance patterns and fit to an Out-of-Africa serial founder effect model, which is known to structure neutral genetic patterns. Moreover, we compare males and females directly, and the true versus false pelvis, in order to examine potential obstetrical effects. Our results indicate evidence for substantial neutral population history effects on pelvic shape variation. They also reveal evidence for the effect of obstetrical constraints, but these affect males and females to equivalent extents. Our results do not deny an important role for selection in regard to specific aspects of human pelvic variation, especially in terms of features associated with body size and proportions. However, our analyses demonstrate that at a global level, the shape of the os coxae reveals substantial evidence for neutral variation. Our analyses thus indicate that population variation in the human pelvis might be used to address important questions concerning population history, just as the human cranium has done.

Human populations across the world vary greatly in cranial morphology. It is highly debated to what extent this variability has accumulated through neutral processes (genetic drift) or through natural selection driven by climate. By taking advantage of recent work showing that geographic distance along landmasses is an excellent proxy for neutral genetic differentiation, we quantify the relative role of drift versus selection in an exceptionally large dataset of human skulls. We show that neutral processes have been much more important than climate in shaping the human cranium. We further demonstrate that a large proportion of the signal for natural selection comes from populations from extremely cold regions. More generally, we show that, if drift is not explicitly accounted for, the effect of natural selection can be greatly overestimated.

One of the main events in the history of our species has been our expansion out of Africa. A clear signature of this expansion has been found on global patterns of neutral genetic variation, whereby a serial founder effect accompanied the colonization of new regions, in turn creating a within-population decrease in neutral genetic diversity with increasing distance from Africa. This same distinctive pattern has also been described for cranial and dental morphological variation in human populations distributed across the globe. Here, we used a data set of postcranial linear measurements for 30 globally distributed human populations, and a climatic data set of minimum annual temperature, maximum annual temperature, and precipitation in order to separate for the first time the relative effect of neutral demographic processes and climatic selection on four long (limb) bones (femur, tibia, radius, and humerus) versus the pelvic bones of the human appendicular skeleton. We implemented a stepwise regression procedure in which phenotypic variance is assumed to be affected by the iterative founder events that accompanied human expansion from Africa, as well as by climate. This model included, as independent factors, geographic distance from central Africa, the three climatic variables, and all possible interactions between the three climatic variables. We excluded all nonsignificant factors by backward stepwise elimination with the aim of identifying the minimal model significantly explaining variation in the phenotypic data. Our results indicate a sharp difference in the way the pelvis and the limb bones reflect the neutral signature of the out-of-Africa expansion. Consistent with previous analyses of the cranium and dentition, pelvic shape variation shows a significant within-population decrease with increasing distance from Africa. However, no such pattern could be found in the long bones. Rather, in the case of both the tibia and the femur, a significant relationship between population-level variance and minimum temperature was demonstrated. Hence, in the case of these limb bones, it is probable that the effects of climatic selection have obliterated the demographic signature of human dispersal from Africa. Our finding that pelvic variation exhibits the neutral effects of demographic history suggests that consideration of this skeletal element might be used to shed light on factors of human population history, just as the cranium has done. KeywordsSerial Founder Effect, Out Of Africa, Anatomically Modern Humans, Pelvis Cover Page FootnoteWe are grateful to Brendon Billings of the Medical School of the University of Witwatersrand and Dr. Ogeto Mwebi of the National Museum of Kenya for allowing access to collections and general assistance. We are also especially grateful to Dr. Benjamin Auerbach for generously making his data freely available. We thank Franz Manni and three anonymous reviewers for constructive comments. This study was funded by Sigma Xi, The Abstract One of the main events in the history of our...

The human birth canal shows a tight fit with the size of the neonate, which can lead to obstetric complications. This is not the case in other apes, and has been explained as the outcome of conflicting evolutionary pressures for bipedal locomotion and parturition of a highly encephalized fetus. Despite the suggested evolutionary constraints on the female pelvis, we show that women are, in fact, extremely variable in the shape of the bony birth canal, with human populations having differently shaped pelvic canals. Neutral evolution through genetic drift and differential migration are largely responsible for the observed pattern of morphological diversity, which correlates well with neutral genetic diversity. Climatic adaptation might have played a role, albeit a minor one, with populations from colder regions showing a more transversally oval shape of the canal inlet. The significant extent of canal shape variation among women from different regions of the world has important implications for modern obstetric practice in multi-ethnic societies, as modern medical understanding has been largely developed on studies of European women.

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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.