How to make a digital reconstruction of the human ribcage.

Up to now, there have been no publication standardizing the digital reconstruction of the modern human ribcage from commingled costo-vertebral material. Consequently, we designed a validated protocol based on anatomical features observed in the literature and the CT scanned ribcages of 10 adult European individuals. After quantifying the shape of these ribcages using 3D geometric morphometrics, we split each vertebra and rib within their corresponding (semi)landmarks. Subsequently, individual bones + (semi)landmarks were imported to LhpFusionBox, commingled and 3D reconstructed. To validate the accuracy of the protocol, we first reconstructed a randomly chosen ribcage three times and then compared these reconstructions to the rest of the sample. Since these reconstructions were closer to their original counterpart than to the others, the remaining sample was reconstructed once. Next, we tested the intra-observer error during reconstructing using the Procrustes distances among the original ribcages and the reconstructions. We observed that first each ribcage reconstruction was clustered to its original counterpart and second there was a learning curve showing an improvement in the reconstruction process over time. Subsequently, we explored general size and shape differences among the original and reconstructed ribcages through a study of centroid size and a permutation test on the Procrustes distances (10,000 permutations), respectively. Specific shape differences between both groups were further examined through a principal component analysis in shape space. None of these analyses found statistical differences between the original and reconstructed ribcages (p > 0.05). Eventually, we extracted the mean shapes of the original ribcages and the reconstructions in order to visualize potential deviations caused by the anatomical considerations of the researcher. These results demonstrate that the protocol is accurate enough to be used when reconstructing a disarticulated human ribcage.

The adaptive evolution of cancer driver genes.

BACKGROUND: Cancer is a life-threatening disease in humans; yet, cancer genes are frequently reported to be under positive selection. This suggests an evolutionary-genetic paradox in which cancer evolves as a secondary product of selection in human beings. However, systematic investigation of the evolution of cancer driver genes is sparse. RESULTS: Using comparative genomics analysis, population genetics analysis and computational molecular evolutionary analysis, the evolution of 568 cancer driver genes of 66 cancer types were evaluated at two levels, selection on the early evolution of humans (long timescale selection in the human lineage during primate evolution, i.e., millions of years), and recent selection in modern human populations (~ 100,000 years). Results showed that eight cancer genes covering 11 cancer types were under positive selection in the human lineage (long timescale selection). And 35 cancer genes covering 47 cancer types were under positive selection in modern human populations (recent selection). Moreover, SNPs associated with thyroid cancer in three thyroid cancer driver genes (CUX1, HERC2 and RGPD3) were under positive selection in East Asian and European populations, consistent with the high incidence of thyroid cancer in these populations. CONCLUSIONS: These findings suggest that cancer can be evolved, in part, as a by-product of adaptive changes in humans. Different SNPs at the same locus can be under different selection pressures in different populations, and thus should be under consideration during precision medicine, especially for targeted medicine in specific populations.

Morphological and morphometric analyses of a late Middle Pleistocene hominin mandible from Hualongdong, China.

Excavations in Hualongdong (HLD), East China, have yielded abundant hominin fossils dated to 300 ka. There is a nearly complete mandible that fits well with a partial cranium, and together they compose the skull labeled as HLD 6. Thus far, detailed morphological description and comparisons of the mandible have not been conducted. Here we present a comprehensive morphological, metric, and geometric morphometric assessment of this mandible and compare it with both adult and immature specimens of Pleistocene hominins and recent modern humans. Results indicate that the HLD 6 mandible exhibits a mosaic morphological pattern characterized by a robust corpus and relatively gracile symphysis and ramus. The moderately developed mental trigone and a clear anterior mandibular incurvation of the HLD 6 mandible are reminiscent of Late Pleistocene hominin and recent modern human morphology. However, the weak expression of all these features indicates that this mandible does not possess a true chin. Moreover, a suite of archaic features that resemble those of Middle Pleistocene hominins includes pronounced alveolar planum, superior transverse torus, thick corpus, a pronounced endocondyloid crest, and a well-developed medial pterygoid tubercle. The geometric morphometric analysis further confirms the mosaic pattern of the HLD 6 mandible. The combination of both archaic and modern human features identified in the HLD 6 mandible is unexpected, given its late Middle Pleistocene age and differs from approximately contemporaneous Homo members such as Xujiayao, Penghu, and Xiahe. This mosaic pattern has never been recorded in late Middle Pleistocene hominin fossil assemblages in East Asia. The HLD 6 mandible provides further support for the high morphological diversity during late Middle Pleistocene hominin evolution. With these findings, it is possible that modern human morphologies are present as early as 300 ka and earlier than the emergence of modern humans in East Asia.

Structural properties of the Late Pleistocene Liujiang femoral diaphyses from southern China.

The characterization of the femoral diaphysis in Pleistocene hominins with chronoecogeographical diversity plays a crucial role in evaluating evolutionary shifts in locomotor behavior and body shape. However, Pleistocene hominin fossil remains in East Asia are scarce and are widely dispersed temporally and spatially, impeding our comprehension of the nature and polarity of morphological trends. Here, we present qualitative and quantitative analyses of the cross-sectional properties and structural organization of diaphyses in two Late Pleistocene hominin femora (Liujiang PA91 and PA92) from southern China, comparing them to other Eurasian and African Pleistocene hominins. By integrating surface features and internal structure, our findings reveal that the Liujiang femora exhibit modern human-like characteristics, including a developed pilaster, a gluteal buttress, and minimum mediolateral breadth located at the midshaft. The presence of a femoral pilaster may relate to posterior cortical reinforcement and an increased anteroposterior bending rigidity along the mid-proximal to mid-distal portion of the diaphysis. Compared to archaic Homo, Liujiang and other Late Pleistocene modern human femora show a thinner mediolateral cortex and lower bending rigidity than the anteroposterior axis, and a lack of medial buttress, potentially indicating functionally related alterations in a range of pelvic and proximal femoral features throughout the Pleistocene. The femoral robusticity of the Liujiang individual resembles that of other Pleistocene hunter-gatherers from East Asia, implying comparable overall mobility or activity levels. The investigation of Liujiang femoral diaphyseal morphology contributes to a more comprehensive understanding of early modern human postcranial structural morphology in East Asia.

Reassessment of the human mandible from Banyoles (Girona, Spain).

Since the discovery of a human mandible in 1887 near the present-day city of Banyoles, northeastern Spain, researchers have generally emphasized its archaic features, including the lack of chin structures, and suggested affinities with the Neandertals or European Middle Pleistocene (Chibanian) specimens. Uranium-series and electron spin resonance dating suggest the mandible dates to the Late Pleistocene (Tarantian), approximately ca. 45-66 ka. In this study, we reassessed the taxonomic affinities of the Banyoles mandible by comparing it to samples of Middle Pleistocene fossils from Africa and Europe, Neandertals, Early and Upper Paleolithic modern humans, and recent modern humans. We evaluated the frequencies and expressions of morphological features and performed a three-dimensional geometric morphometric analysis on a virtual reconstruction of Banyoles to capture overall mandibular shape. Our results revealed no derived Neandertal morphological features in Banyoles. While a principal component analysis based on Euclidean distances from the first two principal components clearly grouped Banyoles with both fossil and recent Homo sapiens individuals, an analysis of the Procrustes residuals demonstrated that Banyoles did not fit into any of the comparative groups. The lack of Neandertal features in Banyoles is surprising considering its Late Pleistocene age. A consideration of the Middle Pleistocene fossil record in Europe and southwest Asia suggests that Banyoles is unlikely to represent a late-surviving Middle Pleistocene population. The lack of chin structures also complicates an assignment to H. sapiens, although early fossil H. sapiens do show somewhat variable development of the chin structures. Thus, Banyoles represents a non-Neandertal Late Pleistocene European individual and highlights the continuing signal of diversity in the hominin fossil record. The present situation makes Banyoles a prime candidate for ancient DNA or proteomic analyses, which may shed additional light on its taxonomic affinities.

The early Aurignacian dispersal of modern humans into westernmost Eurasia.

Documenting the first appearance of modern humans in a given region is key to understanding the dispersal process and the replacement or assimilation of indigenous human populations such as the Neanderthals. The Iberian Peninsula was the last refuge of Neanderthal populations as modern humans advanced across Eurasia. Here we present evidence of an early Aurignacian occupation at Lapa do Picareiro in central Portugal. Diagnostic artifacts were found in a sealed stratigraphic layer dated 41.1 to 38.1 ka cal BP, documenting a modern human presence on the western margin of Iberia ∼5,000 years earlier than previously known. The data indicate a rapid modern human dispersal across southern Europe, reaching the westernmost edge where Neanderthals were thought to persist. The results support the notion of a mosaic process of modern human dispersal and replacement of indigenous Neanderthal populations.

Evolution of Homo in the Middle and Late Pleistocene.

The Middle and Late Pleistocene is arguably the most interesting period in human evolution. This broad period witnessed the evolution of our own lineage, as well as that of our sister taxon, the Neanderthals, and related Denisovans. It is exceptionally rich in both fossil and archaeological remains, and uniquely benefits from insights gained through molecular approaches, such as paleogenetics and paleoproteomics, that are currently not widely applicable in earlier contexts. This wealth of information paints a highly complex picture, often described as 'the Muddle in the Middle,' defying the common adage that 'more evidence is needed' to resolve it. Here we review competing phylogenetic scenarios and the historical and theoretical developments that shaped our approaches to the fossil record, as well as some of the many remaining open questions associated with this period. We propose that advancing our understanding of this critical time requires more than the addition of data and will necessitate a major shift in our conceptual and theoretical framework.

Modern human teeth unearthed from below the ∼128,000-year-old level at Punung, Java: A case highlighting the problem of recent intrusion in cave sediments.

The emergence of modern humans in the eastern edge of the Eurasian Continent is debated between two major models: early (∼130-70 ka) and late (∼50 ka) dispersal models. The former view is grounded mainly on the claims that several cave sites in Southeast Asia and southern China yielded modern human fossils of those early ages, but such reports have been disputed for the lack of direct dating of the human remains and insufficient documentation of stratigraphy and taphonomy. By tracing possible burial process and conducting direct dating for an early Late Pleistocene paleontological site of Punung III, East Java, we here report a case that demonstrates how unexpected intrusion of recent human remains into older stratigraphic levels could occur in cave sediments. This further highlights the need of direct dating and taphonomic assessment before accepting either model. We also emphasize that the state of fossilization of bones and teeth is a useful guide for initial screening of recent intrusion and should be reported particularly when direct dating is unavailable. Additionally, we provide a revised stratigraphy and faunal list of Punung III, a key site that defines the tropical rainforest Punung Fauna during the early Late Pleistocene of the region.

Limits of long-term selection against Neandertal introgression.

Several studies have suggested that introgressed Neandertal DNA was subjected to negative selection in modern humans. A striking observation in support of this is an apparent monotonic decline in Neandertal ancestry observed in modern humans in Europe over the past 45,000 years. Here, we show that this decline is an artifact likely caused by gene flow between modern human populations, which is not taken into account by statistics previously used to estimate Neandertal ancestry. When we apply a statistic that avoids assumptions about modern human demography by taking advantage of two high-coverage Neandertal genomes, we find no evidence for a change in Neandertal ancestry in Europe over the past 45,000 years. We use whole-genome simulations of selection and introgression to investigate a wide range of model parameters and find that negative selection is not expected to cause a significant long-term decline in genome-wide Neandertal ancestry. Nevertheless, these models recapitulate previously observed signals of selection against Neandertal alleles, in particular the depletion of Neandertal ancestry in conserved genomic regions. Surprisingly, we find that this depletion is strongest in regulatory and conserved noncoding regions and in the most conserved portion of protein-coding sequences.

Insights into Modern Human Prehistory Using Ancient Genomes.

The genetic relationship of past modern humans to today's populations and each other was largely unknown until recently, when advances in ancient DNA sequencing allowed for unprecedented analysis of the genomes of these early people. These ancient genomes reveal new insights into human prehistory not always observed studying present-day populations, including greater details on the genetic diversity, population structure, and gene flow that characterized past human populations, particularly in early Eurasia, as well as increased insight on the relationship between archaic and modern humans. Here, we review genetic studies on ∼45000- to 7500-year-old individuals associated with mainly preagricultural cultures found in Eurasia, the Americas, and Africa.

Morphological description and evolutionary significance of 300 ka hominin facial bones from Hualongdong, China.

Late Middle Pleistocene hominins in Africa displaying key modern morphologies by 315 ka are claimed as the earliest Homo sapiens. Evolutionary relationships among East Asian hominins appear complex due to a growing fossil record of late Middle Pleistocene hominins from the region, reflecting mosaic morphologies that contribute to a lack of consensus on when and how the transition to modern humans transpired. Newly discovered 300 ka hominin fossils from Hualongdong, China, provide further evidence to clarify these relationships in the region. In this study, facial morphology of the juvenile partial cranium (HLD 6) is described and qualitatively and quantitatively compared with that of other key Early, Middle, and Late Pleistocene hominins from East Asia, Africa, West Asia, and Europe and with a sample of modern humans. Qualitatively, facial morphology of HLD 6 resembles that of Early and Middle Pleistocene hominins from Zhoukoudian, Nanjing, Dali, and Jinniushan in China, as well as others from Java, Africa, and Europe in some of these features (e.g., supraorbital and malar regions), and Late Pleistocene hominins and modern humans from East Asia, Africa, and Europe in other features (e.g., weak prognathism, flat face and features in nasal and hard plate regions). Comparisons of HLD 6 measurements to group means and multivariate analyses support close affinities of HLD 6 to Late Pleistocene hominins and modern humans. Expression of a mosaic morphological pattern in the HLD 6 facial skeleton further complicates evolutionary interpretations of regional morphological diversity in East Asia. The prevalence of modern features in HLD 6 suggests that the hominin population to which HLD 6 belonged may represent the earliest pre-modern humans in East Asia. Thus, the transition from archaic to modern morphology in East Asian hominins may have occurred at least by 300 ka, which is 80,000 to 100,000 years earlier than previously recognized.

The ripples of modernity: How we can extend paleoanthropology with the extended evolutionary synthesis.

Contemporary understandings of paleoanthropological data illustrate that the search for a line defining, or a specific point designating, "modern human" is problematic. Here we lend support to the argument for the need to look for patterns in the paleoanthropological record that indicate how multiple evolutionary processes intersected to form the human niche, a concept critical to assessing the development and processes involved in the emergence of a contemporary human phenotype. We suggest that incorporating key elements of the Extended Evolutionary Synthesis (EES) into our endeavors offers a better and more integrative toolkit for modeling and assessing the evolution of the genus Homo. To illustrate our points, we highlight how aspects of the genetic exchanges, morphology, and material culture of the later Pleistocene complicate the concept of "modern" human behavior and suggest that multiple evolutionary patterns, processes, and pathways intersected to form the human niche.

The emergence and intensification of early hunter-gatherer niche construction.

Hunter-gatherers, especially Pleistocene examples, are not well-represented in archeological studies of niche construction. However, as the role of humans in shaping environments over long time scales becomes increasingly apparent, it is critical to develop archeological proxies and testable hypotheses about early hunter-gatherer impacts. Modern foragers engage in niche constructive behaviors aimed at maintaining or increasing the productivity of their environments, and these may have had significant ecological consequences over later human evolution. In some cases, they may also represent behaviors unique to modern Homo sapiens. Archeological and paleoenvironmental data show that African hunter-gatherers were niche constructors in diverse environments, which have legacies in how ecosystems function today. These can be conceptualized as behaviorally mediated trophic cascades, and tested using archeological and paleoenvironmental proxies. Thus, large-scale niche construction behavior is possible to identify at deeper time scales, and may be key to understanding the emergence of modern humans.

An early Holocene human skull from Zhaoguo cave, Southwestern China.

OBJECTIVES: In this study, we describe a newly excavated early Holocene human cranium from Guizhou, Southwestern China, namely the Zhaoguo M1 (ZG 1). We aim to evaluate its morphological resemblance with Late Pleistocene human, and Northern and Southern China Neolithic populations. We also aim to infer its position in the process of East Asian population regionalization. MATERIALS AND METHODS: The ZG 1 skull is almost complete, only missing parts of the right parietal and the basicranium around the foramen magnum. Comparative samples include Late Pleistocene humans and Neolithic populations from Northern and Southern China. Univariate and multivariate analyses are carried out in the study. RESULTS: ZG 1 has a dolichocephalic cranium, wide zygomatic breadth, moderate glabella and supraobtial projection, marked canine fossa, and thin cranial vault. The nasal floor, maximum cranial breadth position, and frontal arc proportion are all congruent with modern human. Statistical analysis suggests that ZG 1's measurements are most similar to those of Southern China Neolithic specimens, with some closer to Late Pleistocene humans. CONCLUSION: ZG 1 shows a clear affinity with Southern China Neolithic populations, providing further support that regionalization of morphological variability patterns between Northern and Southern Neolithic populations could have originated at least 10,000 years ago.

Trajectories of cultural innovation from the Middle to Later Stone Age in Eastern Africa: Personal ornaments, bone artifacts, and ocher from Panga ya Saidi, Kenya.

African Middle Stone Age (MSA) populations used pigments, manufactured and wore personal ornaments, made abstract engravings, and produced fully shaped bone tools. However, ongoing research across Africa reveals variability in the emergence of cultural innovations in the MSA and their subsequent development through the Later Stone Age (LSA). When present, it appears that cultural innovations manifest regional variability, suggestive of distinct cultural traditions. In eastern Africa, several Late Pleistocene sites have produced evidence for novel activities, but the chronologies of key behavioral innovations remain unclear. The 3 m deep, well-dated, Panga ya Saidi sequence in eastern Kenya, encompassing 19 layers covering a time span of 78 kyr beginning in late Marine Isotope Stage 5, is the only known African site recording the interplay between cultural and ecological diversity in a coastal forested environment. Excavations have yielded worked and incised bones, ostrich eggshell beads (OES), beads made from seashells, worked and engraved ocher pieces, fragments of coral, and a belemnite fossil. Here, we provide, for the first time, a detailed analysis of this material. This includes a taphonomic, archeozoological, technological, and functional study of bone artifacts; a technological and morphometric analysis of personal ornaments; and a technological and geochemical analysis of ocher pieces. The interpretation of the results stemming from the analysis of OES beads is guided by an ethnoarcheological perspective and field observations. We demonstrate that key cultural innovations on the eastern African coast are evident by 67 ka and exhibit remarkable diversity through the LSA and Iron Age. We suggest the cultural trajectories evident at Panga ya Saidi were shaped by both regional traditions and cultural/demic diffusion.

Mapping carbon and nitrogen isotopic composition of fingernails to demonstrate a rural-urban nutrition transition in the Center-West, Northeast, and Amazon regions of Brazil.

OBJECTIVE: The main objective of this study is to investigate diet patterns among rural and urban populations of the Center-West, Northeast, and Amazon regions of Brazil through the carbon and nitrogen isotopic composition of fingernails, recognizing that the extent of market integration is a key driver of food consumption. MATERIALS AND METHODS: In the Center-West, Northeast, and Amazon regions of Brazil, fingernails were sampled in clusters encompassing a major city, town, and rural village. A total of 2,133 fingernails were analyzed. Fingernails were clipped by donors using fingernail clippers. In the laboratory, samples were cleaned then weighed in small tin capsules before being isotopically analyzed for carbon and nitrogen. RESULTS: The overall mean δ13 C and δ15 N were -19.7 ± 2.8‰ and 10.6 ± 1.1‰, respectively. In the more remote villages, where access to food markets is more challenging, lower δ13 C prevails, suggesting that Brazilian staple foods (rice, beans, and farinha) still dominate. In areas with easier access to food markets, δ13 C values were higher, suggesting a change to a diet based on C4 plants, typical of a Brazilian supermarket diet. The variability among inhabitants in the same location expressed by a significant inverse correlation between δ13 C and δ15 N fingernail values suggested that "market integration" does not affect everyone equally in each community. DISCUSSION AND CONCLUSION: The nutrition transition has not yet reached some remote villages in these regions of Brazil or that the nutrition transition has not yet reached all residents of these remote villages. On the other hand, in several villages there is a considerable adherence to the supermarket diet or that some residents of these villages are already favoring processed food.

Hybridization in human evolution: Insights from other organisms.

During the late Pleistocene, isolated lineages of hominins exchanged genes thus influencing genomic variation in humans in both the past and present. However, the dynamics of this genetic exchange and associated phenotypic consequences through time remain poorly understood. Gene exchange across divergent lineages can result in myriad outcomes arising from these dynamics and the environmental conditions under which it occurs. Here we draw from our collective research across various organisms, illustrating some of the ways in which gene exchange can structure genomic/phenotypic diversity within/among species. We present a range of examples relevant to questions about the evolution of hominins. These examples are not meant to be exhaustive, but rather illustrative of the diverse evolutionary causes/consequences of hybridization, highlighting potential drivers of human evolution in the context of hybridization including: influences on adaptive evolution, climate change, developmental systems, sex-differences in behavior, Haldane's rule and the large X-effect, and transgressive phenotypic variation.

Population structure and the evolution of Homo sapiens in Africa.

It has been proposed that a multiregional model could describe how Homo sapiens evolved in Africa beginning 300,000 years ago. Multiregionalism would require enduring morphological or behavioral differences among African regions and morphological or behavioral continuity within each. African fossils, archeology, and genetics do not comply with either requirement and are unlikely to, because climatic change periodically disrupted continuity and reshuffled populations. As an alternative to multiregionalism, I suggest that reshuffling produced novel gene constellations, including one in which the additive or cumulative effect of newly associated genes enhanced cognitive or communicative potential. Eventual fixation of such a constellation in the lineage leading to modern H. sapiens would explain the abrupt appearance of the African Later Stone Age 50-45 thousand years ago, its nearly simultaneous expansion to Eurasia in the form of the Upper Paleolithic, and the ability of fully modern Upper Paleolithic people to swamp or replace non-modern Eurasians.

The Gravettian child mandible from El Castillo Cave (Puente Viesgo, Cantabria, Spain).

OBJECTIVES: This article documents an incomplete child's mandible found in H. Obermaier's excavation campaign (in 1912) in El Castillo Cave, Spain. This fossil was assigned to what was then considered a phase of the "Aurignacian-delta". MATERIALS AND METHODS: We exhaustively analyzed the original Obermaier documents, with particular attention to those corresponding to the year of the discovery. We extracted a bone sample to radiocarbon date the fossil directly. We also followed established methods to measure, describe and compare the mandible with other human remains. RESULTS: The analysis of Obermaier's documents and new data derived from modern excavations, show that the mandible was discovered in an interior area of the cave. Direct radiocarbon dating yielded a result of 24,720 ± 210 BP and 29,300 - 28,300 cal BP, a date similar to those known for the Gravettian technocomplex both in the El Castillo site and across Europe. The jaw corresponded to a child aged 4-5 years, with modern morphology, but with a certain robustness, especially in the symphyseal region. Comparisons were made with several modern children (Granada, Spitalfields, and Black series) and with immature fossils (European Aurignacian and Gravettian). The few differences between the modern and the fossil children are related to the symphysis and mandibular corpus thickness and height, and to the symphyseal morphology and larger teeth dimensions. Paleoisotopic data for Castillo C correspond with a varied diet. Numerous cutmarks were identified in the midline internal symphyseal region. DISCUSSION AND CONCLUSIONS: The results agree with those published for other fossils of similar age and chronology (e.g., the mandible of the Lagar Velho child) and show clear differences from the jaws of the young Neanderthals. The interpretation of the original data on the mandible discovery may indicate the destruction of a burial and the displacement, by percolation or by a den, at least of part of the skeleton. The perimortem manipulations in the child's mandible are the first described in the Gravettian world of Western Europe.

Cranial measures and ancient DNA both show greater similarity of Neandertals to recent modern Eurasians than to recent modern sub-Saharan Africans.

OBJECTIVES: Ancient DNA analysis has shown that present-day humans of Eurasian ancestry are more similar to Neandertals than are present-day humans of sub-Saharan African ancestry, reflecting interbreeding after modern humans first left Africa. We use craniometric data to test the hypothesis that the crania of recent modern humans show the same pattern. MATERIALS AND METHODS: We computed Mahalanobis squared distances between a published Neandertal centroid based on 37 craniometric traits and each of 2,413 recent modern humans from the Howells global data set (N = 373 sub-Saharan Africans, N = 2,040 individuals of Eurasian descent). RESULTS: The average distance to the Neandertal centroid is significantly lower for Eurasian crania than for sub-Saharan African crania as expected from the findings of ancient DNA (p < 0.001). This result holds when examining distances for separate geographic regions of humans of Eurasian descent (Europeans, Asians, Australasians, Native Americans, and Pacific Islanders). Most of these results are also seen when examining distances partitioning size and shape variation. DISCUSSION: Our results show that the genetic difference in Neandertal ancestry seen in the DNA of present-day sub-Saharan Africans and Eurasians is also found in patterns of recent modern human craniometric variation.