An integrative skeletal and paleogenomic analysis of stature variation suggests relatively reduced health for early European farmers.

Human culture, biology, and health were shaped dramatically by the onset of agriculture ∼12,000 y B.P. This shift is hypothesized to have resulted in increased individual fitness and population growth as evidenced by archaeological and population genomic data alongside a decline in physiological health as inferred from skeletal remains. Here, we consider osteological and ancient DNA data from the same prehistoric individuals to study human stature variation as a proxy for health across a transition to agriculture. Specifically, we compared "predicted" genetic contributions to height from paleogenomic data and "achieved" adult osteological height estimated from long bone measurements for 167 individuals across Europe spanning the Upper Paleolithic to Iron Age (∼38,000 to 2,400 B.P.). We found that individuals from the Neolithic were shorter than expected (given their individual polygenic height scores) by an average of −3.82 cm relative to individuals from the Upper Paleolithic and Mesolithic (P = 0.040) and −2.21 cm shorter relative to post-Neolithic individuals (P = 0.068), with osteological vs. expected stature steadily increasing across the Copper (+1.95 cm relative to the Neolithic), Bronze (+2.70 cm), and Iron (+3.27 cm) Ages. These results were attenuated when we additionally accounted for genome-wide genetic ancestry variation: for example, with Neolithic individuals −2.82 cm shorter than expected on average relative to pre-Neolithic individuals (P = 0.120). We also incorporated observations of paleopathological indicators of nonspecific stress that can persist from childhood to adulthood in skeletal remains into our model. Overall, our work highlights the potential of integrating disparate datasets to explore proxies of health in prehistory.

Morphological integration of the hominoid postcranium.

Previous research has suggested that magnitudes of integration may be distinct in the postcranium of hominoids when compared to other primate species. To test this hypothesis, we estimated and compared magnitudes of integration of eight postcranial bones from three-dimensional surface scans for 57 Hylobates lar, 58 Gorilla gorilla, 60 Pan troglodytes, 60 Homo sapiens, 60 Chlorocebus pygerythrus, and 60 Macaca fascicularis. We tested the hypotheses that 1) magnitudes of integration would be distinct in the postcranium of hominoids compared to cercopithecoids, with the explicit prediction that magnitudes of integration would be lower in hominoids than in cercopithecoids, and 2) girdle elements (scapula, os coxa) would have lower magnitudes of integration across all taxa. Integration was quantified using the integration coefficient of variation from interlandmark distances reflecting anatomical and developmental modules defined according to a priori criteria. A resampling protocol was employed to generate distributions of integration values that were then compared statistically using Mann-Whitney U tests with Bonferroni adjustment. Support for hypothesis 1 was mixed: with the exception of Gorilla, hominoid taxa were less integrated than the cercopithecoids for all anatomical modules. However, Homo, Gorilla, and, to a lesser extent, Pan showed higher integration than Hylobates and the cercopithecoids for homologous limb elements, with magnitudes of integration for both modules being lowest for Hylobates. These results generally support the hypothesis of distinct patterns of magnitudes of integration in the hominoid postcranium. The high integration of Gorilla may be explained by the effects of overall body size. The results supported the predictions of the second hypothesis. Regardless of taxon, the os coxa and scapula were generally the least integrated skeletal elements, while the femur and radius were the most integrated. The lower integration of the girdle elements suggests that the geometric complexities of particular elements may significantly influence study outcomes.

Morphological modularity in the anthropoid axial skeleton.

Previous research has found that hominoids have stronger modularity between limb elements than other anthropoids, suggesting that there is less constraint on morphological diversification (e.g., limb proportions) in hominoids in terms of evolutionary independence. However, degrees of modularity in the axial skeleton have not been investigated across a broad range of anthropoid taxa. Thus, it is unknown whether hominoids also have stronger modularity in the axial skeleton than other anthropoids, which has implications for the evolution of diverse torso morphologies in Miocene apes as well as the evolution of novel characteristics in the skull and vertebrae of fossil hominins. In this study, 12 anthropoid genera were sampled to examine degrees of modularity between axial skeletal elements (i.e., cranium, mandible, vertebrae, and sacrum). Covariance ratio coefficients were calculated using variance/covariance matrices of interlandmark distances for each axial skeletal element to evaluate degrees of modularity. The results showed that Alouatta, Hylobates, Gorilla, Pan, and Homo showed generally stronger modularity than other anthropoid taxa when considering all axial skeletal elements. When only considering the vertebral elements (i.e., vertebrae and sacrum), Alouatta, Hylobates, Gorilla, and Pan showed generally stronger modularity than other anthropoid taxa. Humans showed stronger modularity between the skull and vertebrae than other hominoids. Thus, the evolution of novel characteristics in the skull and vertebral column may have been less constrained in fossil hominins due to the dissociation of trait covariation between axial skeletal elements in hominoid ancestors, thus fostering more evolutionary independence between the skull and vertebral column.

Examination of magnitudes of integration in the catarrhine vertebral column.

The evolution of novel vertebral morphologies observed in humans and other extant hominoids may be related to changes in the magnitudes and/or patterns of covariation among traits. To examine this, we tested magnitudes of integration in the vertebral column of cercopithecoids and hominoids, including humans. Three-dimensional surface scans of 14 vertebral elements from 30 Cercopithecus, 32 Chlorocebus, 39 Macaca, 45 Hylobates, 31 Pan, and 86 Homo specimens were used. A resampling method was used to generate distributions of integration coefficient of variation scores for vertebral elements individually using interlandmark distances. Interspecific comparisons of mean integration coefficient of variation were conducted using Mann-Whitney U tests with Bonferroni adjustment. The results showed that hominoids generally had lower mean integration coefficient of variation than cercopithecoids. In addition, humans showed lower mean integration coefficient of variation than other hominoids in their last thoracic and lumbar vertebrae. Cercopithecoids and Hylobates showed relatively lower mean integration coefficient of variation in cervical vertebrae than in thoracolumbar vertebrae. Pan and Homo showed relatively lower mean integration coefficient of variation in the last thoracic and lumbar vertebrae in the thoracolumbar region, except for the L1 of Pan. The results suggest fewer integration-mediated constraints on the evolution of vertebral morphology in hominoids when compared with cercopithecoids. The weaker magnitudes of integration in lumbar vertebrae in humans when compared with chimpanzees likewise suggest fewer constraints on the evolution of novel lumbar vertebrae morphology in humans.

Ontogenetic changes in magnitudes of integration in the macaque skull.

OBJECTIVES: Magnitudes of morphological integration may constrain or facilitate craniofacial shape variation. The aim of this study was to analyze how the magnitude of integration in the skull of Macaca fascicularis changes throughout ontogeny in relation to developmental and/or functional modules. MATERIALS AND METHODS: Geometric morphometric methods were used to analyze the magnitude of integration in the macaque cranium and mandible in 80 juvenile and 40 adult M. fascicularis specimens. Integration scores in skull modules were calculated using integration coefficient of variation (ICV) of eigenvalues based on a resampling procedure. Resultant ICV scores between the skull as a whole, and developmental and/or functional modules were compared using Mann-Whitney U tests. RESULTS: Results showed that most skull modules were more tightly integrated than the skull as a whole, with the exception of the chondrocranium in juveniles without canines, the chondrocranium/face complex and the mandibular corpus in adults, and the mandibular ramus in all juveniles. The chondrocranium/face and face/mandibular corpus complexes were more tightly integrated in juveniles than adults, possibly reflecting the influences of early brain growth/development, and the changing functional demands of infant suckling and later masticatory loading. This is also supported by the much higher integration of the mandibular ramus in adults compared with juveniles. DISCUSSION: Magnitudes of integration in skull modules reflect developmental/functional mechanisms in M. fascicularis. However, the relationship between "evolutionary flexibility" and developmental/functional mechanisms was not direct or simple, likely because of the complex morphology, multifunctionality, and various ossification origins of the skull.

The relative efficacy of the cranium and os coxa for taxonomic assessment in macaques.

OBJECTIVES: The cranium is generally considered more reliable than the postcranium for assessing primate taxonomy, although recent research suggests that pelvic shape may be equally reliable. However, little research has focused on intrageneric taxonomic discrimination. Here, we test the relative taxonomic efficacy of the cranium and os coxa for differentiating two macaque species, with and without considering sexual dimorphism. MATERIALS AND METHODS: Geometric morphometric analyses were performed on cranial and os coxa landmarks for 119 adult Macaca fascicularis, M. mulatta, and Chlorocebus pygerythrus. Among-group shape variation was examined using canonical variates analyses. Cross-validated discriminant function analysis provided rates of correct group classification. Additionally, average morphological distances were compared with neutral genetic distances. RESULTS: Macaque species were clearly differentiated, both cranially and pelvically, when sex was not considered. Males were more often correctly classified based on the os coxa, while female classification rates were high for both morphologies. Female crania and male os coxa were differentiated approximately the same as genetic distance, while male crania were more similar (convergent), and female os coxa were more divergent than expected based on genetic distance. DISCUSSION: The hypothesis that cranial and os coxal shape can be used to discriminate among macaque species was supported. The cranium was better at differentiating females, while the os coxa was better at differentiating male macaques. Hence, there is no a priori reason for preferring the cranium when assessing intragenetic taxonomic relationships, but the effects of high levels of sexual dimorphism must be corrected for to accurately assess taxonomic signatures.

Biological diversity in an Islamic archaeological population: A radiogenic strontium isotope and craniometric analysis of affinity in Ottoman Romania.

OBJECTIVES: Written accounts, as well as a previous craniometric study, indicate that migrations of non-Europeans and conversions of Europeans to Islam define Ottoman communities in Early Modern Europe. What is less clear are the roles of migration and admixture in generating intra-communal variation. This study combines craniometric with strontium isotope data to compare the cranial affinities of locally born and immigrant individuals. We predict that locally born individuals are more likely than non-locals to show evidence of admixture. MATERIALS AND METHODS: Radiogenic strontium isotope data for 21 Ottomans were compared against archaeological faunal values. Sixteen individuals with intact crania were also measured and compared against two comparative source populations from Anatolia and Europe. Discriminant function analysis assigned unclassified Ottoans to either comparative group based on typicality probabilities, with potential admixture established via intermediate morphology between the two source populations. RESULTS: Strontium isotope values revealed relatively high proportions of non-locals, consistent with high mobility documented historically. The sexes differed, with more males classifying as "typically Anatolian" than females. Locals and non-locals also had different cranial affinity patterns, with most classifying either as "typically Anatolian" or "typically European." Contrary to expectation, none of the locals were identified as intermediate, suggesting admixture rates were relatively low. CONCLUSIONS: Consistent with historical records, the results revealed high levels of extra-regional migration, with most individuals identifiable as either typically Anatolian or European. Moreover, locals and non-locals differed craniometrically, with no signs of admixture between Anatolian migrants and European converts in locals. This suggests intra-communal divisions were maintained.

Multivariate morphometrics, quantitative genetics, and neutral theory: Developing a "modern synthesis" for primate evolutionary morphology.

Anthropologists are increasingly turning to explicit model-bound evolutionary approaches for understanding the morphological diversification of humans and other primate lineages. Such evolutionary morphological analyses rely on three interconnected conceptual frameworks; multivariate morphometrics for quantifying similarity and differences among taxa, quantitative genetics for modeling the inheritance and evolution of morphology, and neutral theory for assessing the likelihood that taxon diversification is due to stochastic processes such as genetic drift. Importantly, neutral theory provides a framework for testing more parsimonious explanations for observed morphological differences before considering more complex adaptive scenarios. However, the consistency with which these concepts are applied varies considerably, which mirrors some of the theoretical obstacles faced during the "modern synthesis" of classical population genetics in the early 20th century. Here, each framework is reviewed and some potential stumbling blocks to the full conceptual integration of multivariate morphometrics, quantitative genetics, and neutral theory are considered.

Diachronic changes in craniofacial morphology among the middle-late Holocene populations from Hehuang region, Northwest China.

OBJECTIVES: This study analyzes craniofacial shape variation in the Hehuang region of Northwest China within a population genetic framework, and takes a diachronic approach to explore the relationship betwee cultural discontinuity and biological continuity/discontinuity in the Hehuang region during the middle to late Holocene. MATERIALS AND METHODS: The sample comprises 76 adult skulls from five archaeological sites, ranging from 4,500 to 1,530 BP. 3D geometric morphometrics, multivariate statistics, quantitative evolutionary genetic and biodistance analyses were performed to study the diachronic variation in craniofacial morphology. Analyses were performed on two cranial modules: the face and the cranial vault, across three major diachronic groups representing the late Neolithic (LNA), the Bronze Age (BA), and the Han-Jin dynasty (HD). RESULTS: Average regional FST for both cranial modules was low, indicating relatively greater variation within diachronic groups than among them. While the LNA and BA groups did not show any significant differences in facial and vault shape, significant craniofacial shape changes were detected between the BA and HD groups. DISCUSSION: The consistent craniofacial morphology during the LNA and the BA, and the shift in morphology between the BA and the HD indicates that cultural discontinuity does not always coincide with biological discontinuity. The Hehuang population evolved in situ with few changes, despite cultural and dietary changes, until the HD when migrations from the Central Plains are associated with extra-local gene flow to the area.

Morphological integration of anatomical, developmental, and functional postcranial modules in the crab-eating macaque (Macaca fascicularis).

OBJECTIVES: Integration and modularity reflect the coordinated action of past evolutionary processes and, in turn, constrain or facilitate phenotypic evolvability. Here, we analyze magnitudes of integration in the macaque postcranium to test whether 20 a priori defined modules are (1) more tightly integrated than random sets of postcranial traits, and (2) are differentiated based on mode of definition, with developmental modules expected to be more integrated than functional or anatomical modules. MATERIALS AND METHODS: The 3D morphometric data collected for eight limb and girdle bones for 60 macaques were collated into anatomical, developmental, and functional modules. A resampling technique was used to create random samples of integration values for each module for statistical comparison. RESULTS: Our results found that not all a priori defined modules were more strongly integrated than random samples of postcranial traits and that specific types of modules did not present consistent patterns of integration. Rather, girdle and joint modules were consistently less integrated than limb modules, and forelimb elements were less integrated than hindlimbs. DISCUSSION: The results suggest that morphometrically complex modules tend to be less integrated than simple limb bones, irrespective of the number of available traits. However, differences in integration of the fore- and hindlimb more likely reflects the multitude of locomotory, feeding, and social functions involved. It remains to be tested whether patterns of integration identified here are primate universals, and to what extent they vary depending on phylogenetic or functional factors.

A craniometric analysis of early modern Romania and Hungary: The roles of migration and conversion in shaping European Ottoman population history.

OBJECTIVES: Debate persists regarding the biological makeup of European Ottoman communities settled during the expansion of the Ottoman Empire during the 16th and 17th centuries, and the roles of conversion and migration in shaping demography and population history. The aim of this study was to perform an assessment of the biological affinities of three European Ottoman series based on craniometric data. MATERIALS AND METHODS: Craniometric data collected from three Ottoman series from Hungary and Romania were compared to European and Anatolian comparative series, selected to represent biological affinity representative of historically recorded migration and conversion influences. Sex-separated samples were analyzed using D2 -matrices, along with principal coordinates and PERMANOVA analyses to investigate biological affinities. Discriminant function analysis was employed to assign Ottoman individuals to two potential classes: European or Anatolian. RESULTS: Affinity analyses show larger than expected biological differences between males and females within each of the Ottoman communities. Discriminant function analyses show that the majority of Ottoman individuals could be classified as either European or Anatolian with a high probability. Moreover, location within Europe proved influential, as the Ottomans from a location of more geopolitical importance (Budapest) diverged from more hinterland communities in terms of biological affinity patterns. DISCUSSION: The results suggest that male and female Ottomans may possess distinct population histories, with males and females divergent from each other in terms of their biological affinities. The Ottoman communities appear diverse in terms of constituting a mix of peoples from different biological backgrounds. The greater distances between sexes from the same community, and the differences between communities, may be evidence that the processes of migration and conversion impacted individual people and groups diversely.

Testing the equivalence of modern human cranial covariance structure: Implications for bioarchaeological applications.

OBJECTIVES: Estimation of the variance-covariance (V/CV) structure of fragmentary bioarchaeological populations requires the use of proxy extant V/CV parameters. However, it is currently unclear whether extant human populations exhibit equivalent V/CV structures. MATERIALS AND METHODS: Random skewers (RS) and hierarchical analyses of common principal components (CPC) were applied to a modern human cranial dataset. Cranial V/CV similarity was assessed globally for samples of individual populations (jackknifed method) and for pairwise population sample contrasts. The results were examined in light of potential explanatory factors for covariance difference, such as geographic region, among-group distance, and sample size. RESULTS: RS analyses showed that population samples exhibited highly correlated multivariate responses to selection, and that differences in RS results were primarily a consequence of differences in sample size. The CPC method yielded mixed results, depending upon the statistical criterion used to evaluate the hierarchy. The hypothesis-testing (step-up) approach was deemed problematic due to sensitivity to low statistical power and elevated Type I errors. In contrast, the model-fitting (lowest AIC) approach suggested that V/CV matrices were proportional and/or shared a large number of CPCs. Pairwise population sample CPC results were correlated with cranial distance, suggesting that population history explains some of the variability in V/CV structure among groups. DISCUSSION: The results indicate that patterns of covariance in human craniometric samples are broadly similar but not identical. These findings have important implications for choosing extant covariance matrices to use as proxy V/CV parameters in evolutionary analyses of past populations.

The relative correspondence of cranial and genetic distances in papionin taxa and the impact of allometric adjustments.

The reconstruction of phylogenetic relationships in the primate fossil record is dependent upon a thorough understanding of the phylogenetic utility of craniodental characters. Here, we test three previously proposed hypotheses for the propensity of primate craniomandibular data to exhibit homoplasy, using a study design based on the relative congruence between cranial distance matrices and a consensus genetic distance matrix ("genetic congruence") for papionin taxa: 1) matrices based on cranial regions subjected to less masticatory strain are more genetically congruent than high-strain cranial matrices; 2) matrices based on cranial regions developing earlier in ontogeny are more genetically congruent than matrices based on regions that develop later; and 3) matrices based on cranial regions with greater anatomical/functional complexity are more genetically congruent than matrices based on anatomically simpler regions. Morphological distance matrices based on the shape of 15 different cranial regions, delineated on the basis of previous catarrhine studies, were statistically compared to a matrix of known genetic distances in papionins. Since sexual dimorphism and allometry are known to characterize this clade, several analytical iterations were conducted: 1) mixed-sex, male-only, and female-only analyses and 2) with and without an allometric scaling adjustment. Across all datasets, the chondrocranium matrix was the most consistently correlated with genetic distances, which is also consistent with previous studies of cercopithecoid taxa; however, there was no support for the internal predictions of the three hypotheses tested. Allometric scaling corrections had the largest impact on the genetic congruence of facial shape matrices, a result consistent with previous studies that have described facial homoplasy in papionin taxa. These findings differ from patterns described for hominoid taxa, suggesting that no single predictive criterion can explain phylogenetic utility of cranial datasets across catarrhine primate taxa. Many of the differences in morphological-genetic matrix correlations could result from different levels of phenotypic integration and evolvability in cercopithecoids and hominoids, suggesting that further study of these phenomena in extant primates is warranted.

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.

Late Pleistocene climate change and the global expansion of anatomically modern 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 hunter-gatherers 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.

A comparison of catarrhine genetic distances with pelvic and cranial morphology: implications for determining hominin phylogeny.

Inferring the evolutionary history of the hominins is necessarily reliant on comparative analyses of fossilized skeletal anatomy. However, the reliability of different primate skeletal regions for recovering phylogenetic relationships is currently poorly understood. Historically, postcranial variation has largely been conceived of as reflecting locomotory and postural adaptation. The shape of the os coxae is central to such discussions given the divergent morphology displayed by the bipedal hominin pelvis relative to other primate taxa. While previous cladistic studies have suggested that postcranial and cranial datasets do not differ in terms of their propensity for homoplasy, methodological issues such as the numbers of characters and their quantification make it difficult to evaluate these findings. Here, we circumvent these problems by constructing morphological distance matrices based on cranial, mandibular and os coxae three-dimensional shape. Statistical comparisons of these morphological distance matrices against a single genetic distance matrix for 11 catarrhine taxa show that cranial and os coxae shape reflect genetic relationships better than the mandible when humans are included, and that the cranium and os coxae do not differ statistically in terms of their genetic correlations. When humans were excluded from the analyses, all three anatomical regions were equally strongly correlated with genetic distance. Moreover, a second analysis focusing solely on os coxae variation of 16 taxa demonstrated that os coxae shape correctly recovers catarrhine taxonomic relationships at the sub-family level, even when humans are included. Taken together, our results suggest that there is no a priori reason to favor cranial shape data over os coxae morphology when inferring the genetic relationships of extant or extinct primate taxa. Morphological similarities between humans and other primates differ depending on the skeletal element, suggesting that combining skeletal elements into a single analysis may provide more accurate reconstructions of genetic relationships.

The interaction of neutral evolutionary processes with climatically-driven adaptive changes in the 3D shape of the human os coxae.

Differences in the breadth of the pelvis among modern human populations and among extinct hominin species have often been interpreted in the light of thermoregulatory adaptation, whereby a larger pelvic girdle would help preserve body temperature in cold environments while a narrower pelvis would help dissipate heat in tropical climates. There is, however, a theoretical problem in interpreting a pattern of variation as evidence of selection without first accounting for the effects of neutral evolutionary processes (i.e., mutation, genetic drift and migration). Here, we analyse 3D configurations of 27 landmarks on the os coxae of 1494 modern human individuals representing 30 male and 23 female populations from five continents and a range of climatic conditions. We test for the effects of climate on the size and shape of the pelvic bone, while explicitly accounting for population history (i.e., geographically-mediated gene flow and genetic drift). We find that neutral processes account for a substantial proportion of shape variance in the human os coxae in both sexes. Beyond the neutral pattern due to population history, temperature is a significant predictor of shape and size variation in the os coxae, at least in males. The effect of climate on the shape of the pelvic bone, however, is comparatively limited, explaining only a small percentage of shape variation in males and females. In accordance with previous hypotheses, the size of the os coxae tends to increase with decreasing temperature, although the significance of the association is reduced when population history is taken into account. In conclusion, the shape and size of the human os coxae reflect both neutral evolutionary processes and climatically-driven adaptive changes. Neutral processes have a substantial effect on pelvic variation, suggesting such factors will need to be taken into account in future studies of human and fossil hominin coxal variation.

Global human mandibular variation reflects differences in agricultural and hunter-gatherer subsistence strategies.

Variation in the masticatory behavior of hunter-gatherer and agricultural populations is hypothesized to be one of the major forces affecting the form of the human mandible. However, this has yet to be analyzed at a global level. Here, the relationship between global mandibular shape variation and subsistence economy is tested, while controlling for the potentially confounding effects of shared population history, geography, and climate. The results demonstrate that the mandible, in contrast to the cranium, significantly reflects subsistence strategy rather than neutral genetic patterns, with hunter-gatherers having consistently longer and narrower mandibles than agriculturalists. These results support notions that a decrease in masticatory stress among agriculturalists causes the mandible to grow and develop differently. This developmental argument also explains why there is often a mismatch between the size of the lower face and the dentition, which, in turn, leads to increased prevalence of dental crowding and malocclusions in modern postindustrial populations. Therefore, these results have important implications for our understanding of human masticatory adaptation.

The microevolution of modern human cranial variation: implications for hominin and primate evolution.

CONTEXT: Reconstructing the evolutionary history of fossil human taxa is heavily reliant on the ability to extract phylogenetic information from patterns of morphological variability. However, attempts to reconstruct phylogenetic relationships from craniodental data in extant primates have yielded inconsistent and inconclusive results. OBJECTIVE: To critically evaluate a recent body of research, conducted within an explicitly quantitative genetics framework, which investigates the extent to which human cranial variation reflects past population history. Possible ways in which to extrapolate these human-specific insights to higher taxonomic levels will also be assessed. RESULTS: A consensus is emerging confirming a largely neutral model for the human cranium, although specific instances of climatic and dietary adaptation have also been uncovered. Also, specific regions of the cranium, delineated according to particular criteria, differ in their relative genetic congruence. However, the genetic congruence patterns identified in modern humans are not replicated in other extant primates, calling their generality into question. CONCLUSIONS: Developing a clearer understanding of the evolution of morphological diversity in extinct taxa requires a different inference approach that focuses on assessing the evolutionary forces that shape these patterns, rather than the identification of particular morphological regions that correlate with genetic relatedness across all primates.