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Structural asymmetries of brain regions associated with lateralised functions have been extensively studied. However, there are fewer morphometric analyses of asymmetries of the gyri and sulci of the entire cortex. The current study assessed cortical asymmetries in a sample of healthy adults (N = 175) from an admixed population from South America. Grey matter volume and surface area of 66 gyri and sulci were quantified on T1 magnetic resonance images. The departure from zero of the differences between left and right hemispheres (L-R), a measure of directional asymmetry (DA), the variance of L-R, and an index of fluctuating asymmetry (FA) were evaluated for each region. Significant departures from perfect symmetry were found for most cortical gyri and sulci. Regions showed leftward asymmetry at the population level in the frontal lobe and superior lateral parts of the parietal lobe. Rightward asymmetry was found in the inferior parietal, occipital, frontopolar, and orbital regions, and the cingulate (anterior, middle, and posterior-ventral). Despite this general pattern, several sulci showed the opposite DA compared to the neighbouring gyri, which remarks the need to consider the neurobiological differences in gyral and sulcal development in the study of structural asymmetries. The results also confirm the absence of DA in most parts of the inferior frontal gyrus and the precentral region. This study contributes with data on populations underrepresented in the databases used in neurosciences. Among its findings, there is agreement with previous results obtained in populations of different ancestry and some discrepancies in the middle frontal and medial parietal regions. A significant DA not reported previously was found for the volume of long and short insular gyri and the central sulcus of the insula, frontomarginal, transverse frontopolar, paracentral, and middle and posterior parts of the cingulate gyrus and sulcus, gyrus rectus, occipital pole, and olfactory sulcus, as well as for the volume and area of the transverse collateral sulcus and suborbital sulcus. Also, several parcels displayed significant variability in the left-right differences, which can be partially attributable to developmental instability, a source of FA. Moreover, a few gyri and sulci displayed ideal FA with non-significant departures from perfect symmetry, such as subcentral and posterior cingulate gyri and sulci, inferior frontal and fusiform gyri, and the calcarine, transverse collateral, precentral, and orbital sulci. Overall, these results show that asymmetries are ubiquitous in the cerebral cortex.

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The morphological changes of the brain and the skull are highly integrated as a result of shared developmental pathways and different types of interactions between them. Shared developmental trajectories between these two structures might be influenced by genetic and environmental factors. Although the effect of environmental factors on neural and craniofacial traits has been extensively studied, less is known about the specific impact of stressful conditions on the coordinated variation between these structures. Here, we test the effect of early nutrient restriction on morphological correspondence between the brain and the endocast. For this purpose, mice exposed to protein or calorie-protein restriction during gestation and lactation were compared with a control group in which dams were fed standard food ad libitum. High-resolution images were obtained after weaning to describe brain and endocranial morphology. By magnetic resonance imaging (MRI), brain volumes were obtained and endocasts were segmented from skull reconstructions derived from micro-computed tomography (microCT). Brain and endocranial volumes were compared to assess the correspondence in size. Shape changes were analyzed using a set of landmarks and semilandmarks on 3D surfaces. Results indicated that brain volume is relatively less affected by undernutrition during development than endocast volume. Shape covariation between the brain and the endocast was found to be quite singular for protein-restricted animals. Procrustes distances were larger between the brain and the endocast of the same specimens than between brains or endocasts of different animals, which means that the greatest similarity is by type of structure and suggests that the use of the endocast as a direct proxy of the brain at this intraspecific scale could have some limitations. In the same line, patterns of brain shape asymmetry were not directly estimated from endocranial surfaces. In sum, our findings indicate that morphological variation and association between the brain and the endocast is modulated by environmental factors and support the idea that head morphogenesis results from complex processes that are sensitive to the pervasive influence of nutrient intake.

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Introduction: The perisylvian region is the cortical core of language and speech. Several accessory sulci have been described in this area, whose presence could modify the results of the automatic quantification of gray matter by popularly used software. This study aimed to assess the expression of accessory sulci in the frontoparietal operculum (FPO) and to evaluate their influence on the gray matter volume estimated by an automatic parcellation of cortical gyri and sulci. Methods: Brain MRI scans of 100 healthy adult volunteers were visually analyzed. The existence of the triangular and diagonal sulci, and the number of accessory sulci in the frontoparietal operculum, were assessed on T1 images. Also, the gray matter volume of gyri and sulci was quantified by an automatized parcellation method. Interhemispheric differences in accessory sulci were evaluated with Chi-square and Wilcoxon paired tests. The effects of the hemisphere, sex, age, total intracranial volume, and accessory sulci on morphometric variables were assessed by linear models. Results: These sulci were found in more than half of the subjects, mostly in the left hemisphere, and showed a significant effect on the gray matter content of the FPO. In particular, the volume of the inferior frontal sulcus, pars opercularis of the inferior frontal gyrus, horizontal ramus of the lateral sulcus, angular gyrus, and postcentral gyrus showed a significant influence on the presence of accessory sulci. Discussion: The prevalence of tertiary sulci in the FPO is high, although their meaning is not yet known. Therefore, they should be considered to reduce the risk of misclassifications of normal variation.

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Brain lateralization is a widespread phenomenon although its expression across primates is still controversial due to the reduced number of species analyzed and the disparity of methods used. To gain insight into the diversification of neuroanatomical asymmetries in non-human primates we analyze the endocasts, as a proxy of external brain morphology, of a large sample of New World monkeys and test the effect of brain size, home range and group sizes in the pattern and magnitude of shape asymmetry. Digital endocasts from 26 species were obtained from MicroCT scans and a set of 3D coordinates was digitized on endocast surfaces. Results indicate that Ateles, Brachyteles, Callicebus and Cacajao tend to have a rightward frontal and a leftward occipital lobe asymmetry, whereas Aotus, Callitrichinae and Cebinae have either the opposite pattern or no directional asymmetry. Such differences in the pattern of asymmetry were associated with group and home range sizes. Conversely, its magnitude was significantly associated with brain size, with larger-brained species showing higher inter-hemispheric differences. These findings support the hypothesis that reduction in inter-hemispheric connectivity in larger brains favors the lateralization and increases the structural asymmetries, whereas the patterns of shape asymmetry might be driven by socio-ecological differences among species.

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The morphology of facial bones is modeled by processes of bone formation and resorption induced by interactions between tissues and compensatory responses. However, the role of remodeling patterns on the morphological changes within and among populations has been scarcely explored. Here, we assess the association between facial shape and the underlying bone cell activity throughout the ontogeny in two Amerindian populations that represent the extremes of craniofacial variation in South America. The sample comprises 71 individuals (36 adults and 35 subadults) representing hunter-gatherers from Patagonia and horticulturists from Northwest Argentina. We analyzed the shape and size of the zygomatic and the maxilla, and compared them with the patterns of bone formation and resorption. Bone formation and resorption were described by quantitative histological analysis of bone surfaces. Morphological changes were described by landmarks and semilandmarks digitized on 3D surfaces obtained from CT images. The results from multivariate statistics analysis show that the patterns of bone remodeling are associated with variation in the morphology of the middle face. We found a similar pattern of facial shape variation along the ontogenetic trajectory in the two samples, and a similar trend in the amount of formation and resorption activities across ages. The main differences between samples were found in the distribution of the areas of bone formation and resorption, possibly associated with mechanical bone response to masticatory loading. These findings provide clues about the processes and mechanisms of bone development involved in the facial morphological differentiation in human populations from southern South America.

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PURPOSE: Brain expansion during ontogeny has been identified as a key factor for explaining the growth pattern of neurocranial bones. However, the dynamics of this relation are only partially understood and a detailed characterization of integrated morphological changes of the brain and the neurocranium along ontogeny is still lacking. The aim of this study was to model the effect of brain growth on cranial bones by means of finite-element analysis (FEA) and geometric morphometric techniques. METHODS: First, we described the postnatal changes in brain size and shape by digitizing coordinates of 3D semilandmarks on cranial endocasts, as a proxy of brain, segmented from CT-scans of an ontogenetic sample. Then, two scenarios of brain growth were simulated: one in which brain volume increases with the same magnitude in all directions, and other that includes the information on the relative expansion of brain regions obtained from morphometric analysis. RESULTS: Results indicate that in the first model, in which a uniform pressure is applied, the largest displacements were localized in the sutures, especially in the anterior and posterior fontanels, as well as the metopic suture. When information of brain relative growth was introduced into the model, displacements were also concentrated in the lambda region although the values along both sides of the neurocranium (parietal and temporal bones) were larger than under the first scenario. CONCLUSION: In sum, we propose a realistic approach to the use of FEA based on morphometric data that offered different results to more simplified models.

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Nutrition is one of the most influential environmental factors affecting the development of different tissues and organs. It is suggested that under nutrient restriction the growth of the brain is spared as a result of the differential allocation of resources from other organs. However, it is not clear whether this sparing occurs brain-wide. Here, we analyzed morphological changes and cell composition in different regions of the offspring mouse brain after maternal exposure to nutrient restriction during pregnancy and lactation. Using high-resolution magnetic resonance imaging, we found that brain regions were differentially sensitive to maternal protein restriction and exhibited particular patterns of volume reduction. The cerebellum was reduced in absolute and relative volume, while cortex volume was relatively preserved. Alterations in cell composition (examined by the isotropic fractionator method) and organization of white matter (measured by diffusor tensor images) were also region specific. These changes were not related to the metabolic rate of the regions and were only partially explained by their specific growth trajectories. This study is a first step towards understanding the mechanisms of regional brain sparing at microstructural and macrostructural levels resulting from undernutrition.

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OBJECTIVES: To assess the intraspecific variation in bone remodeling patterns in modern humans, we studied two populations from southern South America that represent the extremes of morphological variation in this region. We particularly analyzed the ontogenetic changes in the patterns of bone growth remodeling and compared the patterns between samples. MATERIALS AND METHODS: We obtained high-resolution casts of the periosteal surface of the upper and middle face of subadults (n = 36) and adult (n = 36) individuals from a sample of hunter-gatherers from Patagonia and a sample of horticulturists from Northwest Argentina. The areas of bone formation and resorption were registered using an incident-light microscope. We then estimated the average bone remodeling map by sample and age, and performed principal component analysis and multivariate regressions to assess the extension and distribution of these areas across ontogeny and between samples. RESULTS: We found that the remodeling pattern of the glabella, supraorbital arch, frontal process of the maxilla, and a large part of the zygomatic bone is relatively constant in subadults and adults from both sample with a clear predominance of bone formation. In contrast, the middle face is characterized by the spatial alternation between formation and resorption areas, and greater variation with age and between samples. The main differences were found in areas related to chewing and muscle insertions. CONCLUSIONS: Our study provides the first evidence of interpopulation variation in bone growth remodeling and suggests that biomechanical factors can influence the observed patterns. It also underlines the need to account for ecological factors in within and between species comparisons.

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The genetic composition of Amerindian descendants from Patagonia has long been a focus of interest, although the information available is still scarce for many geographic areas. Here, we report the first analysis of the variation in the mitochondrial DNA (mtDNA) control region for an area of northwestern Patagonia, the North of Neuquén, with the aim of studying the processes and historical events that modeled the evolutionary history of these human groups. We analyzed 113 individuals from two localities of northern Neuquén, along with 6 from southern Neuquén and 223 previously published mtDNA sequences from neighboring areas in Argentina and Chile. We estimated the haplotypic variation and spatial structure of molecular variability. Amerindian subhaplogroups predominate in the two samples from northern Neuquén (n = 70), with D1g and C1b13 the most represented, although in different proportions. These samples exhibit Amerindian mtDNA haplotypes similar to the variants from neighboring areas. Most of haplotype variability was within group; variation among groups was relatively low and scarcely associated with geographical space. The most frequent subhaplogroups in northern Neuquén are characteristic of native populations from Patagonia and Chilean Araucanía, and probably originated in the region during the Late Pleistocene or Early Holocene. However, the spatial variation of mtDNA haplotypes departs from a latitudinal pattern and suggests differential levels of gene flow among areas during the Late Holocene, with moderate levels across the North of Neuquén as well as between this area and neighboring populations from Chile, the South of Neuquén, and Río Negro.

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Bone size and shape arise throughout ontogeny as a result of the coordinated activity of osteoblasts and osteoclasts, responsible for bone deposition and resorption, and growth displacements. The modelling processes leave specific microstructural features on the bone surface, which can be used to infer the mechanisms shaping craniofacial traits in extinct and extant species. However, the analysis of bone surfaces from fossils and archaeological samples faces some difficulties related to the bone loss caused by taphonomic factors, and the lack of formal methods for estimating missing information and comparing the patterns of bone modelling among several specimens and samples. The present study provides a new approach for the quantitative analysis of bone formation and resorption patterns obtained from craniofacial surfaces. First, interpolation techniques were used to estimate missing data on high-resolution replicas of the left maxilla in a sample of sub-adult and adult modern humans and sub-adult fossil hominins. The performance of this approach was assessed by simulating variable amounts of missing data. Then, we applied measures of dispersion and central tendency to represent the variation and average pattern of bone modelling within samples. The spatial interpolation resulted in reliable estimations of the type of cell activity (deposition or resorption) in the missing areas, even when large extensions of the bone surface were lost. The quantification of the histological data allowed us to integrate the information of different specimens and depict the areas with higher and lower variation in the bone modelling pattern of the maxilla among specimens. Overall, the main advantages of the quantitative approach used here for generating bone modelling patterns are the high replicability and the possibility of incorporating variation among specimens into the comparisons among samples.

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Mechanisms of resource allocation are essential for maternal and fetal survival, particularly when the availability of nutrients is limited. We investigated the responses of feto-placental development to maternal chronic protein malnutrition to test the hypothesis that maternal low protein diet produces differential growth restriction of placental and fetal tissues, and adaptive changes in the placenta that may mitigate impacts on fetal growth. C57BL/6J female mice were fed either a low-protein diet (6% protein) or control isocaloric diet (20% protein). On embryonic days E10.5, 17.5 and 18.5 tissue samples were prepared for morphometric, histological and quantitative RT-PCR analyses, which included markers of trophoblast cell subtypes. Potential endocrine adaptations were assessed by the expression of Prolactin-related hormone genes. In the low protein group, placenta weight was significantly lower at E10.5, followed by reduction of maternal weight at E17.5, while the fetuses became significantly lighter no earlier than at E18.5. Fetal head at E18.5 in the low protein group, though smaller than controls, was larger than expected for body size. The relative size and shape of the cranial vault and the flexion of the cranial base was affected by E17.5 and more severely by E18.5. The junctional zone, a placenta layer rich in endocrine and energy storing glycogen cells, was smaller in low protein placentas as well as the expression of Pcdh12, a marker of glycogen trophoblast cells. Placental hormone gene Prl3a1 was altered in response to low protein diet: expression was elevated at E17.5 when fetuses were still growing normally, but dropped sharply by E18.5 in parallel with the slowing of fetal growth. This model suggests that nutrients are preferentially allocated to sustain fetal and brain growth and suggests the placenta as a nutrient sensor in early gestation with a role in mitigating impacts of poor maternal nutrition on fetal growth.

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OBJECTIVES: One of the biggest challenges in the study of complex morphologies is to adequately describe shape variation. Here, we assess how the random sampling of surface points automatically obtained performs, when compared with observer-guided sampling procedures, and also evaluate the effect of sliding surface points by bending energy and minimum Procrustes distance. MATERIAL AND METHODS: Three datasets comprising structures with disparate levels of complexity and intrasample variation are as follows: mouse molars, mouse brains, and primate endocasts. Different configurations of 3D coordinates on curves and surfaces were digitized from MRI images and CT scans using semi and fully automated procedures. Shape variables were obtained by Generalized Procrustes Superpositions before and after sliding the pseudolandmarks. Multivariate analyses were used to summarize and compare shape variation. RESULTS: For the primate endocast, the semiautomated and automated strategies yield similar ordinations of specimens. Conversely, the semiautomated strategy better discriminates molar shapes between mouse groups. Shape differences among specimens are not adequately represented by the PCs calculated with surface pseudolandmarks. This is improved when the points are converted into semilandmarks by a sliding criterion. DISCUSSION: Surface semilandmarks automatically obtained from 3D models are promising although they should be used with some caution in complex structures. This approach can be taken as complementary of semiautomated procedures which perform better for assessing shape variation in localized traits previously selected while automated procedures are suitable in studies aimed at comparing overall variation in shape and when there is no previous information about highly variable anatomical regions.

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Native human populations from South America display high levels of craniofacial variation encompassing gracile and robust skulls. Nevertheless, the processes of bone modeling by which morphological variation among populations were attained, remain poorly understood. Here we analyze the relationship between patterns of bone formation and resorption and morphometric variation in the upper face of adults belonging to farmers and hunter-gatherers from northwestern and south Argentina. Our analyses reveal a common pattern of bone modeling of the malar bone characterized by the presence of formation areas. Thus, the larger size and greater development of malar bone exhibited by hunter-gatherers would be linked to a greater magnitude of bone formation activity. Conversely, the glabella and the superciliary arch presented both formation and resorption areas with a variable distribution among individuals. In the extreme corresponding to more robust morphologies, the great development of the glabella is related to the presence of large formation fields, both in the upper region and toward the frontonasal suture. The less robust morphologies show resorption fields at the upper margin of the glabella, which would contribute to the weaker development of this region. The superciliary arch showed a complex relationship between its morphometric and histological variation; the individuals located at both extremes of the shape space presented large resorption areas located on its upper margin. Overall, our results show the existence of intraspecific variation in the patterns of bone modeling in the human upper face.

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Nutritional imbalance is one of the main sources of stress in both extant and extinct human populations. Restricted availability of nutrients is thought to disrupt the buffering mechanisms that contribute to developmental stability and canalization, resulting in increased levels of fluctuating asymmetry (FA) and phenotypic variance among individuals. However, the literature is contradictory in this regard. This study assesses the effect of prenatal nutritional stress on FA and among-individual variance in cranial shape and size using a mouse model of maternal protein restriction. Two sets of landmark coordinates were digitized in three dimensions from skulls of control and protein restricted specimens at E17.5 and E18.5. We found that, by the end of gestation, maternal protein restriction resulted in a significant reduction of skull size. Fluctuating asymmetry in size and shape exceeded the amount of measurement error in all groups, but no significant differences in the magnitude of FA were found between treatments. Conversely, the pattern of shape asymmetry was affected by the environmental perturbation since the angles between the first eigenvectors extracted from the covariance matrix of shape asymmetric component of protein restricted and control groups were not significantly different from the expected for random vectors. In addition, among-individual variance in cranial shape was significantly higher in the protein restricted than the control group at E18.5. Overall, the results obtained from a controlled experiment do not support the view of fluctuating asymmetry of cranial structures as a reliable index for inferring nutritional stress in human populations.

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Previous studies have shown that ecological factors had a significant role in shaping the patterns of craniofacial variation among South American populations. Here, we evaluate whether temperature and diet contributed to facial diversification in small geographic areas. Facial size and shape of 9 osteological samples from central Patagonia (Argentina) were described using 2D landmarks and semilandmarks. Data on mean annual temperature, diet composition (δ(13)C and δ(15)N values) and femoral head maximum breadth, used as a proxy of body mass, were obtained for each sample. We then tested the association of body mass and the ecological variables with facial morphology using spatial regression techniques and a model selection approach. Akaike Information Criterion produced disparate results for both components of facial morphology. The best model for facial size included temperature and body mass proxy, and accounted for more than 80% of variation in size. Lower temperatures were related to larger facial sizes. Body mass was negatively associated with facial size and showed no relationship with the temperature. This suggests a relatively independent variation of cranial traits and body mass at the spatial scale studied here. Facial shape was not associated with the temperature or diet composition, contrasting with the patterns observed at larger spatial scales. Our results point out that the effect of climatic variables on cranial traits might be a source of morphological differentiation not only at large scales but also in small geographic areas, and that size and shape display a differential preservation of environmental signals.

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Organism size is controlled by interactions between genetic and environmental factors mediated by hormones with systemic and local effects. As changes in size are usually not isometric, a considerable diversity in shape can be generated through modifications in the patterns of ontogenetic allometry. In this study we evaluated the role of timing and dose of growth hormone (GH) release on growth and correlated shape changes in craniofacial bones. Using a longitudinal study design, we analyzed GH deficient mice treated with GH supplementation commencing pre- and post-puberty. We obtained 3D in vivo micro-CT images of the skull between 21 and 60 days of age and used geometric morphometrics to analyze size and shape changes among control and GH deficient treated and non-treated mice. The variable levels of circulating GH altered the size and shape of the adult skull, and influenced the cranial base, vault, and face differently. While cranial base synchondroses and facial sutures were susceptible to either the direct or indirect effect of GH supplementation, its effect was negligible on the vault. Such different responses support the role of intrinsic growth trajectories of skeletal components in controlling the modifications induced by systemic factors. Contrary to the expected, the timing of GH treatment did not have an effect on catch-up growth. GH levels also altered the ontogenetic trajectories by inducing changes in their location and extension in the shape space, indicating that differences arose before 21 days and were further accentuated by a truncation of the ontogenetic trajectories in GHD groups.

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Modifications of ontogenetic allometries play an important role in patterning the shape differentiation among populations. This study evaluates the influence of size variation on craniofacial shape disparity among human populations from South America and assesses whether the morphological disparity observed at the interpopulation level resulted from a variable extension of the same ontogenetic allometry, or whether it arose as a result of divergences in the pattern of size-related shape changes. The size and shape of 282 adult and subadult crania were described by geometric morphometric-based techniques. Multivariate regressions were used to evaluate the influence of size on shape differentiation between and within populations, and phylogenetic comparative methods were used to take into account the shared evolutionary history among populations. The phylogenetic generalized least-squares models showed that size accounts for a significant amount of shape variation among populations for the vault and face but not for the base, suggesting that the three modules did not exhibit a uniform response to changes in overall growth. The common slope test indicated that patterns of evolutionary and ontogenetic allometry for the vault and face were similar and characterized by a heightening of the face and a lengthening of the vault with increasing size. The conservation of the same pattern of shape changes with size suggests that differences in the extent of growth contributed to the interpopulation cranial shape variation and that certain directions of morphological change were favored by the trait covariation along ontogeny.

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Environmental perturbations of many kinds influence growth and development. Little is known, however, about the influence of environmental factors on the patterns of phenotypic integration observed in complex morphological traits. We analyze the changes in phenotypic variance-covariance structure of the rat skull throughout the early postnatal ontogeny (from birth to weaning) and evaluate the effect of intrauterine growth retardation (IUGR) on this structure. Using 2D coordinates taken from lateral radiographs obtained every 4 days, from birth to 21 days old, we show that the pattern of covariance is temporally dynamic from birth to 21 days. The environmental perturbation provoked during pregnancy altered the skull growth, and reduced the mean size of the IUGR group. These environmental effects persisted throughout lactancy, when the mothers of both groups received a standard diet. More strikingly, the effect grew larger beyond this point. Altering environmental conditions did not affect all traits equally, as revealed by the low correlations between covariance matrices of treatments at the same age. Finally, we found that the IUGR treatment increased morphological integration as measured by the scaled variance of eigenvalues. This increase coincided and is likely related to an increase in morphological variance in this group. This result is expected if somatic growth is a major determinant of covariance structure of the skull. In summary, our findings suggest that environmental perturbations experienced in early ontogeny alter fundamental developmental processes and are an important factor in shaping the variance-covariance structure of complex phenotypic traits.

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The purpose of this article is to examine the patterns of evolutionary relationships between human populations from the later Late Holocene (1,500-100 years BP) of southern South America on the basis of dental morphometric data. We tested the hypotheses that the variation observed in this region would be explained by the existence of populations with different phylogenetic origin or differential action of gene flow and genetic drift. In this study, we analyzed permanent teeth from 17 samples of male and female adult individuals from throughout southern South America. We measured mesiodistal and buccolingual diameters at the base of the crown, along the cement-enamel junction. The results of multiple regression analysis and a mantel correlogram indicate the existence of spatial structure in dental shape variation, as the D(2) Mahalanobis distance between samples increases with increasing geographical distance between them. In addition, the correlation test results show a trend toward reduction of the internal variation of samples with increasing latitude. The detected pattern of dental variation agrees with the one expected as an outcome of founder serial effects related to an expansion of range during the initial occupation of southern South America.

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The knowledge of processes involved in morphological variation requires the integrated analysis of evolutionary and ecological factors. Here, we investigate the factors responsible for dental variation among human populations from southern South America. The aim of this work is to test the correspondence of dental size and shape variation with geographical, molecular (i.e. mtDNA) and ecological (i.e. climate, diet and food preparation) variables employing comparative phylogenetic methods, which have not previously been extensively applied at a within-species level. The results of the Procrustes analysis show a significant association of shape variables with molecular distance and geography, whereas dental size is not associated with molecular or geographical distances among groups. Phylogenetic generalized least-squares analysis, which takes into account the evolutionary autocorrelation among populations, shows a significant relationship between dental size variation and diet, while temperature and pottery do not correspond with dental size or shape. Specifically, groups with diets rich in carbohydrates, as well as the maritime hunter-gatherers, have the smallest teeth. In summary, our results support ecological factors as the dominant factor on dental size diversification in this region, while evolutionary relationships account for variation in dental shape.

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