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Fossils and artifacts from Herto, Ethiopia, include the most complete child and adult crania of early

In humans, obstetrical difficulties arise from the large head and broad shoulders of the neonate relative to the maternal birth canal. Various characteristics of human cranial development, such as the relatively small head of neonates compared with adults and the delayed fusion of the metopic suture, have been suggested to reflect developmental adaptations to obstetrical constraints. On the other hand, it remains unknown whether the shoulders of humans also exhibit developmental features reflecting obstetrical adaptation. Here we address this question by tracking the development of shoulder width from fetal to adult stages in humans, chimpanzees, and Japanese macaques. Compared with nonhuman primates, shoulder development in humans follows a different trajectory, exhibiting reduced growth relative to trunk length before birth and enhanced growth after birth. This indicates that the perinatal developmental characteristics of the shoulders likely evolved to ease obstetrical difficulties such as shoulder dystocia in humans., ヒトは小さく生まれて大きく育つ --その秘密は鎖骨にあり--. 京都大学プレスリリース. 2022-04-13.

The dispersal of modern humans from Africa is now well documented with genetic data that track population history, as well as gene flow between populations. Phenetic skeletal data, such as cranial and pelvic morphologies, also exhibit a dispersal-from-Africa signal, which, however, tends to be blurred by the effects of local adaptation and in vivo phenotypic plasticity, and that is often deteriorated by postmortem damage to skeletal remains. These complexities raise the question of which skeletal structures most effectively track neutral population history. The cavity system of the inner ear (the so-called bony labyrinth) is a good candidate structure for such analyses. It is already fully formed by birth, which minimizes postnatal phenotypic plasticity, and it is generally well preserved in archaeological samples. Here we use morphometric data of the bony labyrinth to show that it is a surprisingly good marker of the global dispersal of modern humans from Africa. Labyrinthine morphology tracks genetic distances and geography in accordance with an isolation-by-distance model with dispersal from Africa. Our data further indicate that the neutral-like pattern of variation is compatible with stabilizing selection on labyrinth morphology. Given the increasingly important role of the petrous bone for ancient DNA recovery from archaeological specimens, we encourage researchers to acquire 3D morphological data of the inner ear structures before any invasive sampling. Such data will constitute an important archive of phenotypic variation in present and past populations, and will permit individual-based genotype-phenotype comparisons.

Significance Human brains differ substantially from those of great apes, and equally important differences exist between their braincases. However, it remains unclear to which extent evolutionary changes in brain structure are related to changes in braincase structure. To study this question, we use combined computed tomography (CT) and MRI head data of humans and chimpanzees and quantify the spatial correlations between brain sulci and cranial sutures. We show that the human brain–braincase relationships are unique compared to chimpanzees and other great apes and that structural rearrangements in the brain and in the braincase emerged independently during human evolution. These data serve as an important frame of reference to identify and quantify evolutionary changes in brain and braincase structures in fossil hominin endocasts., Throughout hominin evolution, the brain of our ancestors underwent a 3-fold increase in size and substantial structural reorganization. However, inferring brain reorganization from fossil hominin neurocrania (=braincases) remains a challenge, above all because comparative data relating brain to neurocranial structures in living humans and great apes are still scarce. Here we use MRI and same-subject spatially aligned computed tomography (CT) and MRI data of humans and chimpanzees to quantify the spatial relationships between these structures, both within and across species. Results indicate that evolutionary changes in brain and neurocranial structures are largely independent of each other. The brains of humans compared to chimpanzees exhibit a characteristic posterior shift of the inferior pre- and postcentral gyri, indicative of reorganization of the frontal opercular region. Changes in human neurocranial structure do not reflect cortical reorganization. Rather, they reflect constraints related to increased encephalization and obligate bipedalism, resulting in relative enlargement of the parietal bones and anterior displacement of the cerebellar fossa. This implies that the relative position and size of neurocranial bones, as well as overall endocranial shape (e.g., globularity), should not be used to make inferences about evolutionary changes in the relative size or reorganization of adjacent cortical regions of fossil hominins.

The brains of modern humans differ from those of great apes in size, shape, and cortical organization, notably in frontal lobe areas involved in complex cognitive tasks, such as social cognition, tool use, and language. When these differences arose during human evolution is a question of ongoing debate. Here, we show that the brains of early

Objectives The bony labyrinth of the inner ear has special relevance when tracking phenotypic evolution because it is often well preserved in fossil and modern primates. Here we track the evolution of the bony labyrinth of anthropoid primates during the Mio−Plio−Pleistocene—the time period that gave rise to the extant great apes and humans. Materials and Methods We use geometric morphometrics to analyze labyrinthine morphology in a wide range of extant and fossil anthropoids, including New World and Old World monkeys, apes, and humans; fossil taxa are represented by Aegyptopithecus, Microcolobus, Epipliopithecus, Nacholapithecus, Oreopithecus, Ardipithecus, Australopithecus, and Homo. Results Our results show that the morphology of the anthropoid bony labyrinth conveys a statistically significant phylogenetic signal especially at the family level. The bony labyrinthine morphology of anthropoids is also in part associated with size, but does not cluster by locomotor adaptations. The Miocene apes examined here, regardless of inferred locomotor behaviors, show labyrinthine morphologies distinct from modern great apes. Discussion Our results suggest that labyrinthine variation contains mixed signals and alternative explanations need to be explored, such as random genetic drift and neutral phenotypic evolution, as well as developmental constraints. The observed pattern in fossil and extant hominoids also suggests that an additional factor, for example, prenatal brain development, could have potentially had a larger role in the evolutionary modification of the bony labyrinth than hitherto recognized.

© The Author(s) 2018., The present study attempted to reconstruct 3D brain shape of Neanderthals and early Homo sapiens based on computational neuroanatomy. We found that early Homo sapiens had relatively larger cerebellar hemispheres but a smaller occipital region in the cerebrum than Neanderthals long before the time that Neanderthals disappeared. Further, using behavioural and structural imaging data of living humans, the abilities such as cognitive flexibility, attention, the language processing, episodic and working memory capacity were positively correlated with size-adjusted cerebellar volume. As the cerebellar hemispheres are structured as a large array of uniform neural modules, a larger cerebellum may possess a larger capacity for cognitive information processing. Such a neuroanatomical difference in the cerebellum may have caused important differences in cognitive and social abilities between the two species and might have contributed to the replacement of Neanderthals by early Homo sapiens., This study was supported by a Grant-in-Aid for Scientifc Research on Innovative Areas “Replacement of Neanderthals by Modern Humans: Testing Evolutionary Models of Learning” from the Japanese Ministry of Education, Culture, Sports, Science and Technology to T.A. (#22101001), N.O. (#22101006), and H.C.T. (#22101007) respectively, and by Swiss NSF grant 31003A_135470 to C.P.E.Z., CGL-2015-63648-P MINECO, Spain to M.B., and Te Calleva Foundation and the Human Origins Research Fund to C.S.

Inferring the morphology of the last common ancestor of humans, chimpanzees and gorillas is a matter of ongoing debate. Recent findings and reassessment of fossil hominins leads to the hypothesis that the last common ancestor was not extant African ape-like. However, an African great-ape-like ancestor with knuckle walking features still remains plausible and the most parsimonious scenario. Here we address this question via an evolutionary developmental approach, comparing taxon-specific patterns of shape change of the femoral diaphysis from birth to adulthood in great apes, humans, and macaques. While chimpanzees and gorillas exhibit similar locomotor behaviors, our data provide evidence for distinct ontogenetic trajectories, indicating independent evolutionary histories of femoral ontogeny. Our data further indicate that anthropoid primates share a basic pattern of femoral diaphyseal ontogeny that reflects shared developmental constraints. Humans escaped from these constraints via differential elongation of femur., ヒトの祖先はチンパンジーやゴリラには似ていない --発生パターンの比較から二足歩行の起源に迫る--. 京都大学プレスリリース. 2018-02-01.

Because brains do not fossilize, the internal surface of the braincase (endocast) serves as an important source of information about brain growth, development, and evolution. Recent studies of endocranial morphology and development in great apes, fossil hominins, and modern humans have revealed taxon-specific differences. However, it remains to be investigated to which extent differences in endocranial morphology reflect differences in actual brain morphology and development, and to which extent they reflect different interactions of the brain and its case with the cranial base and face. Here we address this question by analyzing the effects of cranial integration on endocranial morphology. We test the 'spatial packing' and 'facial orientation' hypotheses, which propose that size and orientation of the neurocranium relative to the viscerocranium influence endocranial shape. Results show that a substantial proportion of endocranial shape variation along and across ontogenetic trajectories is due to cranial integration. Specifically, the uniquely globular shape of the human endocast mainly results from the combination of an exceptionally large brain with a comparatively small face. Overall, thus, cranial integration has pervasive effects on endocranial morphology, and only a comparatively small proportion of inter- and intra-taxon variation can directly be associated with variation in brain morphology.

We describe a general Godunov-type splitting for numerical simulations of the Fisher–Kolmogorov–Petrovski–Piskunov growth and diffusion equation on a world map with Neumann boundary conditions. The procedure is semi-implicit, hence quite stable. Our principal application for this solver is modeling human population dispersal over geographical maps with changing paleovegetation and paleoclimate in the late Pleistocene. As a proxy for carrying capacity we use Net Primary Productivity (NPP) to predict times for human arrival in the Americas., PLoS ONE, 12 (1), ISSN:1932-6203

Individual-based models (IBMs) of human populations capture spatio-temporal dynamics using rules that govern the birth, behavior, and death of individuals. We explore a stochastic IBM of logistic growth-diffusion with constant time steps and independent, simultaneous actions of birth, death, and movement that approaches the Fisher-Kolmogorov model in the continuum limit. This model is well-suited to parallelization on high-performance computers. We explore its emergent properties with analytical approximations and numerical simulations in parameter ranges relevant to human population dynamics and ecology, and reproduce continuous-time results in the limit of small transition probabilities. Our model prediction indicates that the population density and dispersal speed are affected by fluctuations in the number of individuals. The discrete-time model displays novel properties owing to the binomial character of the fluctuations: in certain regimes of the growth model, a decrease in time step size drives the system away from the continuum limit. These effects are especially important at local population sizes of, PLoS ONE, 12 (4), ISSN:1932-6203

The multiple skeletal components of the skull originate asynchronously and their developmental schedule varies across amniotes. Here we present the embryonic ossification sequence of 134 species, covering all major groups of mammals and their close relatives. This comprehensive data set allows reconstruction of the heterochronic and modular evolution of the skull and the condition of the last common ancestor of mammals. We show that the mode of ossification (dermal or endochondral) unites bones into integrated evolutionary modules of heterochronic changes and imposes evolutionary constraints on cranial heterochrony. However, some skull-roof bones, such as the supraoccipital, exhibit evolutionary degrees of freedom in these constraints. Ossification timing of the neurocranium was considerably accelerated during the origin of mammals. Furthermore, association between developmental timing of the supraoccipital and brain size was identified among amniotes. We argue that cranial heterochrony in mammals has occurred in concert with encephalization but within a conserved modular organization., The skeletal components of the skull develop at different times in mammals. Here, Koyabu et al. show that the mode of bone ossification constrains the timing of bone formation and find an association between the developmental timing of the supraoccipital bone and brain size.

In our study on human pelvic development (1), we propose two hypotheses explaining the narrowing of the female pelvis during postreproductive life. Hypothesis 1 proposes ultimate, evolutionary causes, reflecting selective pressures acting on postreproductive life. Hypothesis 2 proposes proximate, developmental causes, reflecting reduced estrogen levels during postreproductive life. Mitteroecker and Fischer (2) replicate one part of our study—that concerning adult pelvic development—with an independent dataset, taking into account age effects on pelvic shape variation not considered in their earlier study (3). (Note that the smoothing functions used in our study only serve for visual guidance, not for statistical testing.) Rather than testing our hypotheses (1) with new … [↵][1]1To whom correspondence should be addressed. Email: marcia{at}aim.uzh.ch. [1]: #xref-corresp-1-1

Several well-known Miocene catarrhines, including Proconsul heseloni, have been inferred to combine quadrupedal walking in an arboreal substrate with a significant amount of climbing during locomotion. The degree to which some of these species were adapted to perform these behaviors is not fully understood due to the mosaic of 'ape-like' and 'monkey-like' traits identified in the forelimb. Given these unique combinations of forelimb features in the fossils, we report on forelimb traits that should be emphasized when investigating skeletal adaptation to quadrupedalism (defined in this manuscript as symmetrical gait movement on horizontal supports, excluding knuckle-walking) and climbing (including both vertical climbing and clambering). We investigate the correspondence between: 1) quadrupedalism and two well-known forelimb traits, humeral torsion and olecranon process length, and 2) climbing and phalangeal curvature. We also test the degree of phylogenetic signal in these relationships using phylogenetic generalized least-squares and branch length transformation methods in order to determine the models of best-fit. We present models that can be used to predict proportions of quadrupedalism and climbing in extant and extinct anthropoid taxa. Each trait-behavior correlation is significant and characterized by an absence of phylogenetic signal. Thus, we employ models assuming a star phylogeny to predict locomotor proportions. The climbing model based on phalangeal curvature and a proxy for size provides the most accurate predictions of behavior across anthropoids. The two quadrupedalism models are less accurate, but distinguish highly quadrupedal species from those that are not. Predictive equations based on these traits support the inference that P. heseloni performed a high proportion of quadrupedalism with a significant climbing component. The degree of phalangeal curvature measured in Pliopithecus vindobonensis predicts that this Miocene catarrhine species performed a proportion of climbing similar to Proconsul, while humeral torsion and olecranon process length provide conflicting inferences of quadrupedal locomotion in this species.

The proximal femoral morphology of fossil hominins is routinely interpreted in terms of muscular topography and associated locomotor modes. However, the detailed correspondence between hard and soft tissue structures in the proximal femoral region of extant great apes is relatively unknown, because dissection protocols typically do not comprise in-depth osteological descriptions. Here, we use computed tomography and virtopsy (virtual dissection) for non-invasive examination of the femoropelvic musculoskeletal anatomy in Pan troglodytes, P. paniscus, Gorilla gorilla, Pongo pygmaeus, and Homo sapiens. Specifically, we analyze the topographic relationship between muscle attachment sites and surface structures of the proximal femoral shaft such as the lateral spiral pilaster. Our results show that the origin of the vastus lateralis muscle is anterior to the insertion of gluteus maximus in all examined great ape specimens and humans. In gorillas and orangutans, the insertion of gluteus maximus is on the inferior (anterolateral) side of the lateral spiral pilaster. In chimpanzees, however, the maximus insertion is on its superior (posteromedial) side, similar to the situation in modern humans. These findings support the hypothesis that chimpanzees and humans exhibit a shared-derived musculoskeletal topography of the proximal femoral region, irrespective of their different locomotor modes, whereas gorillas and orangutans represent the primitive condition. Caution is thus warranted when inferring locomotor behavior from the surface topography of the proximal femur of fossil hominins, as the morphology of this region may contain a strong phyletic signal that tends to blur locomotor adaptation.

Since the beginnings of paleoanthropology, immature fossil hominin specimens have marked important but highly contested cornerstones of research. Long deemed as not representative of a fossil species' morphology, immature hominins are now in the center of scientific attention, and an increasing interest in evolutionary developmental questions has made developmental paleoanthropology a vibrant field of research. Here we report on recent advances in this field, which result from a combination of new methods to reconstruct fossil ontogenies with insights from evo-devo research on extant species.

The interpretation of patterns of cranial pneumatization in terms of evolution, development, and function is controversial, because these structures exhibit extreme diversity and variability among and within taxa. However, there is general consensus that air-filled spaces are formed by invasion of mucous epithelial tissue from the nasopharyngeal cavity into the surrounding cranial bones. This investigation presents a morphogenetic model of pneumatization, which combines empirical data about epithelial growth with physical concepts of surface growth. The study develops a model that defines growth equations with a minimum number of system parameters to simulate the invasion of mucous tissue and air-filled spaces into the cancellous compartment of cranial bones. Computer simulations show that tuning a small set of model parameters permits generation of a wide diversity of morphologies mimicking natural air-filled spaces. Comparison of virtual with actual morphologies yields new insights into possible factors controlling the process of cranial pneumatization.

From birth to adulthood, the human brain expands by a factor of 3.3, compared with 2.5 in chimpanzees [DeSilva J and Lesnik J (2006) Chimpanzee neonatal brain size: Implications for brain growth in Homo erectus. J Hum Evol 51: 207–212]. How the required extra amount of human brain growth is achieved and what its implications are for human life history and cognitive development are still a matter of debate. Likewise, because comparative fossil evidence is scarce, when and how the modern human pattern of brain growth arose during evolution is largely unknown. Virtual reconstructions of a Neanderthal neonate from Mezmaiskaya Cave (Russia) and of two Neanderthal infant skeletons from Dederiyeh Cave (Syria) now provide new comparative insights: Neanderthal brain size at birth was similar to that in recent Homo sapiens and most likely subject to similar obstetric constraints. Neanderthal brain growth rates during early infancy were higher, however. This pattern of growth resulted in larger adult brain sizes but not in earlier completion of brain growth. Because large brains growing at high rates require large, late-maturing, mothers [Leigh SR and Blomquist GE (2007) in Campbell CJ et al. Primates in perspective; pp 396–407], it is likely that Neanderthal life history was similarly slow, or even slower-paced, than in recent H. sapiens .

The Plio-Pleistocene site of Dmanisi, Georgia, has yielded a rich fossil and archaeological record documenting an early presence of the genus Homo outside Africa. Although the craniomandibular morphology of early Homo is well known as a result of finds from Dmanisi and African localities, data about its postcranial morphology are still relatively scarce. Here we describe newly excavated postcranial material from Dmanisi comprising a partial skeleton of an adolescent individual, associated with skull D2700/D2735, and the remains from three adult individuals. This material shows that the postcranial anatomy of the Dmanisi hominins has a surprising mosaic of primitive and derived features. The primitive features include a small body size, a low encephalization quotient and absence of humeral torsion; the derived features include modern-human-like body proportions and lower limb morphology indicative of the capability for long-distance travel. Thus, the earliest known hominins to have lived outside of Africa in the temperate zones of Eurasia did not yet display the full set of derived skeletal features.

Between 2003 and 2005, the Peştera cu Oase, Romania yielded a largely complete early modern human cranium, Oase 2, scattered on the surface of a Late Pleistocene hydraulically displaced bone bed containing principally the remains of Ursus spelaeus . Multiple lines of evidence indicate an age of ≈40.5 thousand calendar years before the present (≈35 ka 14 C B.P.). Morphological comparison of the adolescent Oase 2 cranium to relevant Late Pleistocene human samples documents a suite of derived modern human and/or non-Neandertal features, including absence of a supraorbital torus, subrectangular orbits, prominent canine fossae, narrow nasal aperture, level nasal floor, angled and anteriorly oriented zygomatic bones, a high neurocranium with prominent parietal bosses and marked sagittal parietal curvature, superiorly positioned temporal zygomatic root, vertical auditory porous, laterally bulbous mastoid processes, superiorly positioned posterior semicircular canal, absence of a nuchal torus and a suprainiac fossa, and a small occipital bun. However, these features are associated with an exceptionally flat frontal arc, a moderately large juxtamastoid eminence, extremely large molars that become progressively larger distally, complex occlusal morphology of the upper third molar, and relatively anteriorly positioned zygomatic arches. Moreover, the featureless occipital region and small mastoid process are at variance with the large facial skeleton and dentition. This unusual mosaic in Oase 2, some of which is paralleled in the Oase 1 mandible, indicates both complex population dynamics as modern humans dispersed into Europe and significant ongoing human evolution once modern humans were established within Europe.

Due to the scarcity of the fossil record, in vivo changes in the dentognathic system of early Homo are typically documented at the level of individual fossil specimens, and it remains difficult to draw population-level inferences about dietary habits, diet-related activities and lifestyle from individual patterns of dentognathic alterations. The Plio-Pleistocene hominin sample from Dmanisi (Georgia), dated to 1.77 million years ago, offers a unique opportunity to study in vivo changes in the dentognathic system of individuals belonging to a single paleodeme of early Homo.We analyze dentognathic pathologies in the Dmanisi sample, and in comparative samples of modern Australian and Greenlander hunter-gatherer populations, applying clinical protocols of dentognathic diagnostics.The Dmanisi hominins exhibit a similarly wide diversity and similar incidence of dentognathic pathologies as the modern human hunter-gatherer population samples investigated here. Dmanisi differs from the modern population samples in several respects: At young age tooth wear is already advanced, and pathologies are more prevalent. At old age, hypercementosis is substantial.Results indicate that dentognathic pathologies and disease trajectories are largely similar in early Homo and modern humans, but that the disease load was higher in early Homo, probably as an effect of higher overall stress on the dentognathic system. Am J Phys Anthropol 160:229-253, 2016. © 2016 Wiley Periodicals, Inc.

How and when the Americas were populated remains contentious. Using ancient and modern genome-wide data, we find that the ancestors of all present-day Native Americans, including Athabascans and Amerindians, entered the Americas as a single migration wave from Siberia no earlier than 23 thousand years ago (KYA), and after no more than 8,000-year isolation period in Beringia. Following their arrival to the Americas, ancestral Native Americans diversified into two basal genetic branches around 13 KYA, one that is now dispersed across North and South America and the other is restricted to North America. Subsequent gene flow resulted in some Native Americans sharing ancestry with present-day East Asians (including Siberians) and, more distantly, Australo-Melanesians. Putative ‘Paleoamerican’ relict populations, including the historical Mexican Pericúes and South American Fuego-Patagonians, are not directly related to modern Australo-Melanesians as suggested by the Paleoamerican Model.

Kennewick Man, referred to as the Ancient One by Native Americans, is a male human skeleton discovered in Washington state (USA) in 1996 and initially radiocarbon dated to 8,340-9,200 calibrated years before present (BP). His population affinities have been the subject of scientific debate and legal controversy. Based on an initial study of cranial morphology it was asserted that Kennewick Man was neither Native American nor closely related to the claimant Plateau tribes of the Pacific Northwest, who claimed ancestral relationship and requested repatriation under the Native American Graves Protection and Repatriation Act (NAGPRA). The morphological analysis was important to judicial decisions that Kennewick Man was not Native American and that therefore NAGPRA did not apply. Instead of repatriation, additional studies of the remains were permitted. Subsequent craniometric analysis affirmed Kennewick Man to be more closely related to circumpacific groups such as the Ainu and Polynesians than he is to modern Native Americans. In order to resolve Kennewick Man's ancestry and affiliations, we have sequenced his genome to ∼1× coverage and compared it to worldwide genomic data including for the Ainu and Polynesians. We find that Kennewick Man is closer to modern Native Americans than to any other population worldwide. Among the Native American groups for whom genome-wide data are available for comparison, several seem to be descended from a population closely related to that of Kennewick Man, including the Confederated Tribes of the Colville Reservation (Colville), one of the five tribes claiming Kennewick Man. We revisit the cranial analyses and find that, as opposed to genome-wide comparisons, it is not possible on that basis to affiliate Kennewick Man to specific contemporary groups. We therefore conclude based on genetic comparisons that Kennewick Man shows continuity with Native North Americans over at least the last eight millennia.

Newly discovered Homo remains, stone artifacts, and animal fossils from Dmanisi, Republic of Georgia, provide a basis for better understanding patterns of hominin evolution and behavior in Eurasia ca. 1.77 million years ago. Here we describe a fourth skull that is nearly complete, lacking all but one of its teeth at the time of death. Both the maxillae and the mandible exhibit extensive bone loss due to resorption. This individual is similar to others from the site but supplies information about variation in brain size and craniofacial anatomy within the Dmanisi paleodeme. Although this assemblage presents numerous primitive characters, the Dmanisi skulls are best accommodated within the species H. erectus. On anatomical grounds, it is argued that the relatively small-brained and lightly built Dmanisi hominins may be ancestral to African and Far Eastern branches of H. erectus showing more derived morphology.

Summary While the braincase of adult Neanderthals had a similar volume to that of modern humans from the same period, differences in endocranial shape suggest that brain morphology differed between modern humans and Neanderthals. When and how these differences arose during evolution and development is a topic of ongoing research, with potential implications for species-specific differences in brain and cognitive development, and in life history [1,2]. Earlier research suggested that Neanderthals followed an ancestral mode of brain development, similar to that of our closest living relatives, the chimpanzees [2–4]. Modern humans, by contrast, were suggested to follow a uniquely derived mode of brain development just after birth, giving rise to the characteristically globular shape of the adult human brain case [2,4,5]. Here, we re-examine this hypothesis using an extended sample of Neanderthal infants. We document endocranial development during the decisive first two years of postnatal life. The new data indicate that Neanderthals followed largely similar modes of endocranial development to modern humans. These findings challenge the notion that human brain and cognitive development after birth is uniquely derived [2,4].

The St. Césaire 1 Neanderthal skeleton of a young adult individual is unique in its association with Châtelperronian artifacts from a level dated to ca. 36,000 years ago. Computer-tomographic imaging and computer-assisted reconstruction of the skull revealed a healed fracture in the cranial vault. When paleopathological and forensic diagnostic standards are applied, the bony scar bears direct evidence for the impact of a sharp implement, which was presumably directed toward the individual during an act of interpersonal violence. These findings add to the evidence that Neanderthals used implements not only for hunting and food processing, but also in other behavioral contexts. It is hypothesized that the high intra-group damage potential inherent to weapons might have represented a major factor during the evolution of hominid social behavior.

Schwartz et al . hold that variation among the Dmanisi skulls reflects taxic diversity. The morphological observations to support their hypothesis, however, are partly incorrect, and not calibrated against intraspecific variation in living taxa. After proper adjustment, Schwartz et al .’s data are fully compatible with the hypothesis of a single paleodeme of early Homo at Dmanisi.

Homo neanderthalensis has a unique combination of craniofacial features that are distinct from fossil and extant ‘anatomically modern’ Homo sapiens (modern humans). Morphological evidence, direct isotopic dates1 and fossil mitochondrial DNA from three Neanderthals2,3 indicate that the Neanderthals were a separate evolutionary lineage for at least 500,000 yr. However, it is unknown when and how Neanderthal craniofacial autapomorphies (unique, derived characters) emerged during ontogeny. Here we use computerized fossil reconstruction4 and geometric morphometrics5,6 to show that characteristic differences in cranial and mandibular shape between Neanderthals and modern humans arose very early during development, possibly prenatally, and were maintained throughout postnatal ontogeny. Postnatal differences in cranial ontogeny between the two taxa are characterized primarily by heterochronic modifications of a common spatial pattern of development. Evidence for early ontogenetic divergence together with evolutionary stasis of taxon-specific patterns of ontogeny is consistent with separation of Neanderthals and modern humans at the species level.

The purpose of this study was to assess the importance of stereolithographic models (SLMs) for preoperative diagnosis and planning in craniofacial surgery and to examine whether these models offer valuable additional information as compared to normal CT scans and 3D CT images. Craniofacial SLMs of 20 patients with craniomaxillofacial pathology were made. A helical volume CT scan of the anatomic area involved delivered the necessary data for their construction. These were built with an SLA 250 stereolithography apparatus (3D-Systems, Valencia, CA, USA), steered by FORM-IT/DCS software (University of Zurich, Switzerland). The stereolithography models were classified according to pathology, type of surgery and their relevance for surgical planning. Though not objectively measurable, it was beyond doubt that relevant additional information for the surgeon was obtained in cases of hypertelorism, severe asymmetries of the neuro- and viscerocranium, complex cranial synostoses and large skull defects. The value of these models as realistic "duplicates" of complex or rare dysmorphic craniofacial pathology for the purpose of creating a didactic collection should also be emphasized. The models proved to be less useful in cases of consolidated fractures of the periorbital and naso-ethmoidal complex, except where there was major dislocation.

The Plio-Pleistocene hominin sample from Dmanisi (Georgia), dated to 1.77 million years ago, is unique in offering detailed insights into patterns of morphological variation within a paleodeme of early Homo. Cranial and dentoalveolar morphologies exhibit a high degree of diversity, but the causes of variation are still relatively unexplored. Here we show that wear-related dentoalveolar remodeling is one of the principal mechanisms causing mandibular shape variation in fossil Homo and in modern human hunter–gatherer populations. We identify a consistent pattern of mandibular morphological alteration, suggesting that dental wear and compensatory remodeling mechanisms remained fairly constant throughout the evolution of the genus Homo. With increasing occlusal and interproximal tooth wear, the teeth continue to erupt, the posterior dentition tends to drift in a mesial direction, and the front teeth become more upright. The resulting changes in dentognathic size and shape are substantial and need to be taken into account in comparative taxonomic analyses of isolated hominin mandibles. Our data further show that excessive tooth wear eventually leads to a breakdown of the normal remodeling mechanisms, resulting in dentognathic pathologies, tooth loss, and loss of masticatory function. Complete breakdown of dentognathic homeostasis, however, is unlikely to have limited the life span of early Homo because this effect was likely mediated by the preparation of soft foods.

BackgroundAcquisition of bipedality is a hallmark of human evolution. How bipedality evolved from great ape-like locomotor behaviors, however, is still highly debated. This is mainly because it is difficult to infer locomotor function, and even more so locomotor kinematics, from fossil hominin long bones. Structure-function relationships are complex, as long bone morphology reflects phyletic history, developmental programs, and loading history during an individual's lifetime. Here we discriminate between these factors by investigating the morphology of long bones in fetal and neonate great apes and humans, before the onset of locomotion.Methodology/principal findingsComparative morphometric analysis of the femoral diaphysis indicates that its morphology reflects phyletic relationships between hominoid taxa to a greater extent than taxon-specific locomotor adaptations. Diaphyseal morphology in humans and chimpanzees exhibits several shared-derived features, despite substantial differences in locomotor adaptations. Orangutan and gorilla morphologies are largely similar, and likely represent the primitive hominoid state.Conclusions/significanceThese findings are compatible with two possible evolutionary scenarios. Diaphyseal morphology may reflect retained adaptive traits of ancestral taxa, hence human-chimpanzee shared-derived features may be indicative of the locomotor behavior of our last common ancestor. Alternatively, diaphyseal morphology might reflect evolution by genetic drift (neutral evolution) rather than selection, and might thus be more informative about phyletic relationships between taxa than about locomotor adaptations. Both scenarios are consistent with the hypothesis that knuckle-walking in chimpanzees and gorillas resulted from convergent evolution, and that the evolution of human bipedality is unrelated to extant great ape locomotor specializations.

Hominin evolution is characterized by two main trends, transition to bipedality and increase in brain size. Fossil evidence shows that both trends had a major impact on the structure and function of the hominin skull. This chapter asks how evolutionary modification of the cranial ontogenetic program led to morphological reorganization of the hominin skull and ultimately to hominin cranial diversity. Three major mechanisms of evolutionary developmental reorganization are proposed: modified prenatal development of the cranial base and face reflects adaptation to bipedality; high rates of neurocranial growth during early postnatal ontogeny are essential to attain large brain sizes; taxon-specific modification of facial development reflects dietary adaptation and-in the genus Homo-a general trend toward neoteny.

Entheseal changes are traditionally included in a large array of skeletal features commonly referred to as ‘‘skeletal markers of activity.’’ However, medical studies and recent anthropological analyses of identified skeletal series suggest a complex combination of physiological and biomechanical factors underlying the variability of such ‘‘markers.’’ The aim of this study is to examine the relationship between age, sex, physical activity, and entheseal variability. To this end, 23 postcranial entheses are examined in a large (N 5 484) Italian contemporary skeletal series using standardized scoring methods. The sample comprises subjects of known age, sex and, mostly, occupation. Results show a strong relationship between age and entheseal changes. Differences between sexes are also highlighted, while the effects of physical activity appear moderate. Altogether, our study indicates that entheseal morphology primarily reflects the age of an individual, while correlation with lifetime activity remains ambiguous.

Interosseous sutures exhibit highly variable patterns of interdigitation and corrugation. Recent research has identified fundamental molecular mechanisms of suture formation, and computer models have been used to simulate suture morphogenesis. However, the role of bone strain in the development of complex sutures is largely unknown, and measuring suture morphologies beyond the evaluation of fractal dimensions remains a challenge. Here we propose a morphogenetic model of suture formation, which is based on the paradigm of Laplacian interface growth. Computer simulations of suture morphogenesis under various boundary conditions generate a wide variety of synthetic sutural forms. Their morphologies are quantified with a combination of Fourier analysis and principal components analysis, and compared with natural morphological variation in an ontogenetic sample of human interparietal suture lines. Morphometric analyses indicate that natural sutural shapes exhibit a complex distribution in morphospace. The distribution of synthetic sutures closely matches the natural distribution. In both natural and synthetic systems, sutural complexity increases during morphogenesis. Exploration of the parameter space of the simulation system indicates that variation in strain and/or morphogen sensitivity and viscosity of sutural tissue may be key factors in generating the large variability of natural suture complexity.

The Dmanisi hominins inhabited a northern temperate habitat in the southern Caucasus, approximately 1.8 million years ago. This is the oldest population of hominins known outside of Africa. Understanding the set of anatomical and behavioral traits that equipped this population to exploit their seasonal habitat successfully may shed light on the selection pressures shaping early members of the genus Homo and the ecological strategies that permitted the expansion of their range outside of the African subtropics. The abundant stone tools at the site, as well as taphonomic evidence for butchery, suggest that the Dmanisi hominins were active hunters or scavengers. In this study, we examine the locomotor mechanics of the Dmanisi hind limb to test the hypothesis that the inclusion of meat in the diet is associated with an increase in walking and running economy and endurance. Using comparative data from modern humans, chimpanzees, and gorillas, as well as other fossil hominins, we show that the Dmanisi hind limb was functionally similar to modern humans, with a longitudinal plantar arch, increased limb length, and human-like ankle morphology. Other aspects of the foot, specifically metatarsal morphology and tibial torsion, are less derived and similar to earlier hominins. These results are consistent with hypotheses linking hunting and scavenging to improved walking and running performance in early Homo. Primitive retentions in the Dmanisi foot suggest that locomotor evolution continued through the early Pleistocene.

The recent reconstruction of the Sahelanthropus tchadensis cranium (TM 266-01-60-1) provides an opportunity to examine in detail differences in cranial shape between this earliest-known hominid, African apes, and other hominid taxa. Here we compare the reconstruction of TM 266-01-60-1 with crania of African apes, humans, and several Pliocene hominids. The results not only confirm that TM 266-01-60-1 is a hominid but also reveal a unique mosaic of characters. The TM 266-01-60-1 reconstruction shares many primitive features with chimpanzees but overall is most similar to Australopithecus , particularly in the basicranium. However, TM 266-01-60-1 is distinctive in having the combination of a short subnasal region associated with a vertical upper face that projects substantially in front of the neurocranium. Further research is needed to determine the evolutionary relationships between Sahelanthropus and the known Miocene and Pliocene hominids.

Another hominid skull has been recovered at Dmanisi (Republic of Georgia) from the same strata in which hominid remains have been reported previously. The Dmanisi site dated to ∼1.75 million years ago has now produced craniofacial portions of several hominid individuals, along with many well-preserved animal fossils and quantities of stone artifacts. Although there are certain anatomical differences among the Dmanisi specimens, the hominids do not clearly represent more than one taxon. We assign the new skull provisionally to Homo erectus (= ergaster ). The Dmanisi specimens are the most primitive and small-brained fossils to be grouped with this species or any taxon linked unequivocally with genus Homo and also the ones most similar to the presumed habilis -like stem. We suggest that the ancestors of the Dmanisi population dispersed from Africa before the emergence of humans identified broadly with the H. erectus grade.

The geometric morphometric analysis of shape variation in complex biological structures such as the human skull poses a number of specific challenges: the registration of homologous morphologies, the treatment of bilateral symmetry, the graphical representation of form variability in three dimensions and the interpretation of the results in terms of differential growth processes. To visualize complex patterns of shape change, we propose an alternative to classical Cartesian deformation grids in the style of D'Arcy W. Thompson. Reference to the surface structures of the organism under investigation permits a comprehensive visual grasp of shape change and its tentative interpretation in terms of differential growth. The application of this method to the analysis of human craniofacial shape variation reveals distinct modes of growth and development of the neurocranial and viscerocranial regions of the skull. Our data further indicate that variations in the orientation of the viscerocranium relative to the neurocranium impinge on the shapes of the face and the cranial vault.

Studies comparing phenotypic variation with neutral genetic variation in modern humans have shown that genetic drift is a main factor of evolutionary diversification among populations. The genetic population history of our closest living relatives, the chimpanzees and bonobos, is now equally well documented, but phenotypic variation among these taxa remains relatively unexplored, and phenotype-genotype correlations are not yet documented. Also, while the adult phenotype is typically used as a reference, it remains to be investigated how phenotype-genotye correlations change during development. Here we address these questions by analyzing phenotypic evolutionary and developmental diversification in the species and subspecies of the genus Pan. Our analyses focus on the morphology of the femoral diaphysis, which represents a functionally constrained element of the locomotor system. Results show that during infancy phenotypic distances between taxa are largely congruent with non-coding (neutral) genotypic distances. Later during ontogeny, however, phenotypic distances deviate from genotypic distances, mainly as an effect of heterochronic shifts between taxon-specific developmental programs. Early phenotypic differences between Pan taxa are thus likely brought about by genetic drift while late differences reflect taxon-specific adaptations.

Since 1980 an operation which reestablishes the blood outflow from occluded hepatic veins was performed in 7 patients with Budd-Chiari syndrome by one of us (A. Senning). Using extracorporeal circulation a dorsocranial cylindrical resection of the liver including the confluence of the occluded hepatic veins was performed by transcaval approach. The incised right atrium was sutured around the resected liver area. There was one intraoperative death. In 6 patients with a mean postoperative follow-up of 19.2 months (4-42 months), the patency of hepatoatrial anastomosis was documented by angiography or Doppler-2d-echocardiography. Four patients are free of symptoms and signs of Budd-Chiari syndrome. In one of two patients with associated cirrhosis compression of inferior vena cava reoccurred and in another patient esophageal varices persist. We conclude, that the hepatoatrial anastomosis is an effective treatment of Budd-Chiari syndrome.