Sarah E. Freidline

Middle Pleistocene human facial morphology in an evolutionary and developmental context.

Neanderthals and modern humans exhibit distinct facial architectures. The patterning of facial morphology of their predecessors, the Middle Pleistocene humans, is more mosaic showing a mix of archaic and modern morphologies. Significant changes in facial size and robusticity occurred throughout Pleistocene human evolution, resulting in temporal trends in both facial reduction and enlargement. However, the allometric patterning in facial morphology in archaic humans is not well understood. This study explores temporal trends in facial morphology in order to gain a clearer understanding of the polarity of features, and describes the allometric patterning of facial shape. The modern human sample comprises cross-sectional growth series of four morphologically distinct human populations. The fossil sample covers specimens from the Middle Pleistocene to the Upper Paleolithic. We digitized landmarks and semilandmarks on surface and computed tomography scans and analyzed the Procrustes shape coordinates. Principal component analyses were performed, and Procrustes distances were used to identify phenetic similarities between fossil hominins. In order to explore the influence of size on facial features, allometric trajectories were calculated for fossil and modern human groups, and developmental simulations were performed. We show that facial features can be used to separate Pleistocene humans into temporal clusters. The distinctly modern human pattern of facial morphology is already present around 170 ka. Species- and population-specific facial features develop before two years of age, and several of the large-scale facial differences between Neanderthals and Middle Pleistocene humans are due to scaling along a shared allometric trajectory. These features include aspects of the frontal bone, browridge morphology, nasal aperture size and facial prognathism. Infraorbital surface topography and orientation of the midface in the European Middle Pleistocene hominins is intermediate between the African Middle Pleistocene and Neanderthal condition. This could suggest that the European Middle Pleistocene hominins display incipient Neanderthal features.

Ontogenetic and static allometry in the human face: contrasting Khoisan and Inuit.

OBJECTIVES Regional differences in modern human facial features are present at birth, and ontogenetic allometry contributes to variation in adults. However, details regarding differential rates of growth and timing among regional groups are lacking. We explore ontogenetic and static allometry in a cross-sectional sample spanning Africa, Europe and North America, and evaluate tempo and mode in two regional groups with very different adult facial morphology, the Khoisan and Inuit. MATERIALS AND METHODS Semilandmark geometric morphometric methods, multivariate statistics and growth simulations were used to quantify and compare patterns of facial growth and development. RESULTS Regional-specific facial morphology develops early in ontogeny. The Inuit has the most distinct morphology and exhibits heterochronic differences in development compared to other regional groups. Allometric patterns differ during early postnatal development, when significant increases in size are coupled with large amounts of shape changes. All regional groups share a common adult static allometric trajectory, which can be attributed to sexual dimorphism, and the corresponding allometric shape changes resemble developmental patterns during later ontogeny. DISCUSSION The amount and pattern of growth and development may not be shared between regional groups, indicating that a certain degree of flexibility is allowed for in order to achieve adult size. In early postnatal development the face is less constrained compared to other parts of the cranium allowing for greater evolvability. The early development of region-specific facial features combined with heterochronic differences in timing or rate of growth, reflected in differences in facial size, suggest different patterns of postnatal growth.

Bone growth dynamics of the facial skeleton and mandible in Gorilla gorilla and Pan troglodytes

Adult craniofacial morphology results from complex processes that involve growth by bone modelling and interactions of skeletal components to keep a functional and structural balance. Previous analyses of growth dynamics in humans revealed critical changes during late ontogeny explaining particular morphological features in our species. Data on bone modelling patterns from other primate species could help us to determine whether postnatal changes in the growth dynamics of the craniofacial complex are human specific or are shared with other primates. However, characterizations of bone modelling patterns through ontogeny in non-human hominids are scarce and restricted to isolated data on facial and mandibular regions. In the present study, we analyse the bone modelling patterns in an ontogenetic series of Pan and Gorilla to infer the growth dynamics of their craniofacial complex during postnatal development. Our results show that both Pan troglodytes and Gorilla gorilla are characterized by species-specific bone modelling patterns indicative of a mainly forward growth direction during postnatal development. Both species show minor but consistent ontogenetic changes in the distribution of bone modelling fields in specific regions of the face and mandible, in contrast to other regions which show more constant bone modelling patterns. In addition, we carry out a preliminary integrative study merging histological and geometric morphometric data. Both approaches yield highly complementary data, each analysis providing details on specific growth dynamics unavailable to the other. Moreover, geometric morphometric data show that ontogenetic variation in the modelling pattern of the mandibular ramus may be linked to sexual dimorphism.

3D landmark and semilandmark geometric morphometric analysis of the Zuttiyeh fronto-zygomatic fragment

Cortisol, as a biomarker of the generalized adaptive stress response, can provide critical information on the physiological effects of behavior. However, group-living animals face multiple interacting stressors from their social and ecological environments. While recent research has revealed the impact of particular social stressors in isolation, few studies have examined how diverse factors contribute to long-term stress hormone variation. We applied multivariate analyses to a 10-year dataset to investigate urinary cortisol variation in wild female chimpanzees (Pan troglodytes schweinfurthii) in Kibale National Park, Uganda. In interindividual contrasts, older females had higher cortisol levels, as did females that were low-ranking for their age. Over time, cortisol was significantly predicted by rates of aggression, particularly female-targeted aggression. The effect of male-female aggression was most pronounced for cycling, swollen females, who are the most frequent targets of sexual coercion. On the other hand, lactating females were strongly affected by group size, such that cortisol levels were elevated when parties contained more females and fewer males. Fruit consumption was an important covariate for lactating females, who experienced higher cortisol when the diet was poor. Our data indicate that both social and energetic factors contribute to stress variation in female chimpanzees, but that energetic stress increases in significance for females facing high reproductive costs. Our study also contributes to growing evidence that direct and indirect competition, while subtle in their expression, can have substantial impacts on female chimpanzees.

Author Correction: New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens

Fossil evidence points to an African origin of Homo sapiens from a group called either H. heidelbergensis or H. rhodesiensis. However, the exact place and time of emergence of H. sapiens remain obscure because the fossil record is scarce and the chronological age of many key specimens remains uncertain. In particular, it is unclear whether the present day ‘modern’ morphology rapidly emerged approximately 200 thousand years ago (ka) among earlier representatives of H. sapiens or evolved gradually over the last 400 thousand years. Here we report newly discovered human fossils from Jebel Irhoud, Morocco, and interpret the affinities of the hominins from this site with other archaic and recent human groups. We identified a mosaic of features including facial, mandibular and dental morphology that aligns the Jebel Irhoud material with early or recent anatomically modern humans and more primitive neurocranial and endocranial morphology. In combination with an age of 315 ± 34 thousand years (as determined by thermoluminescence dating), this evidence makes Jebel Irhoud the oldest and richest African Middle Stone Age hominin site that documents early stages of the H. sapiens clade in which key features of modern morphology were established. Furthermore, it shows that the evolutionary processes behind the emergence of H. sapiens involved the whole African continent.

Middle Pleistocene human facial morphology

Supplement: Program of the 81st Annual Meeting of the American Association of Physical Anthropologists (2012)The catacomb of Saints Peter and Marcellinus located in the south-east of Rome approximately contains 25 000 graves dated from the 3rd to the 5th century AD. Seven newly discovered rooms having an unusual organization in the heart of the catacomb were investigated in 2003. Excavations of these rooms revealed a mass grave, where 3000 corpses were laid together. These individuals were stacked in rows apparently following a common fatal incident. Presumably, this epidemic crisis occurred between the 1st and the 3rd century AD. The specific funerary treatment (textile wrapping and plaster) recalls mummification and might be related to exogenous practices, possibly connected to Early Christians. Moreover, the presence of rare and expensive materials (e.g., Baltic amber, resins and gold threads) may indicate a high social status. Stable isotope analyses (carbon, nitrogen and oxygen) of bone collagen, bone apatite and tooth hydroxyapatite were carried out on 111 individuals to obtain further information on their diet and residential mobility. Additionally, a study of dental nonmetric traits was conducted on 200 individuals to define the biological distance between the deceased and to assess their phenetic similarity. The combination of these two approaches will bring new insight into the homogeneity of the Early population of the catacomb of Saints Peter and Marcellinus and the relationship between funerary practices and geographical origin of buried individuals.