Mandibular ramus shape variation and ontogeny in Homo sapiens and Homo neanderthalensis.

As the interface between the mandible and cranium, the mandibular ramus is functionally significant and its morphology has been suggested to be informative for taxonomic and phylogenetic analyses. In primates, and particularly in great apes and humans, ramus morphology is highly variable, especially in the shape of the coronoid process and the relationship of the ramus to the alveolar margin. Here we compare ramus shape variation through ontogeny in Homo neanderthalensis to that of modern and fossil Homo sapiens using geometric morphometric analyses of two-dimensional semilandmarks and univariate measurements of ramus angulation and relative coronoid and condyle height. Results suggest that ramus, especially coronoid, morphology varies within and among subadult and adult modern human populations, with the Alaskan Inuit being particularly distinct. We also identify significant differences in overall anterosuperior ramus and coronoid shapes between H. sapiens and H. neanderthalensis, both in adults and throughout ontogeny. These shape differences are subtle, however, and we therefore suggest caution when using ramus morphology to diagnose group membership for individual specimens of these taxa. Furthermore, we argue that these morphologies are unlikely to be representative of differences in masticatory biomechanics and/or paramasticatory behaviors between Neanderthals and modern humans, as has been suggested by previous authors. Assessments of ontogenetic patterns of shape change reveal that the typical Neanderthal ramus morphology is established early in ontogeny, and there is little evidence for divergent postnatal ontogenetic allometric trajectories between Neanderthals and modern humans as a whole. This analysis informs our understanding of intraspecific patterns of mandibular shape variation and ontogeny in H. sapiens and can shed further light on overall developmental and life history differences between H. sapiens and H. neanderthalensis.

Craniomandibular form and body size variation of first generation mouse hybrids: A model for hominin hybridization.

Hybridization occurs in a number of mammalian lineages, including among primate taxa. Analyses of ancient genomes have shown that hybridization between our lineage and other archaic hominins in Eurasia occurred numerous times in the past. However, we still have limited empirical data on what a hybrid skeleton looks like, or how to spot patterns of hybridization among fossils for which there are no genetic data. Here we use experimental mouse models to supplement previous studies of primates. We characterize size and shape variation in the cranium and mandible of three wild-derived inbred mouse strains and their first generation (F1) hybrids. The three parent taxa in our analysis represent lineages that diverged over approximately the same period as the human/Neanderthal/Denisovan lineages and their hybrids are variably successful in the wild. Comparisons of body size, as quantified by long bone measurements, are also presented to determine whether the identified phenotypic effects of hybridization are localized to the cranium or represent overall body size changes. The results indicate that hybrid cranial and mandibular sizes, as well as limb length, exceed that of the parent taxa in all cases. All three F1 hybrid crosses display similar patterns of size and form variation. These results are generally consistent with earlier studies on primates and other mammals, suggesting that the effects of hybridization may be similar across very different scenarios of hybridization, including different levels of hybrid fitness. This paper serves to supplement previous studies aimed at identifying F1 hybrids in the fossil record and to introduce further research that will explore hybrid morphologies using mice as a proxy for better understanding hybridization in the hominin fossil record.

Mandibular ramus shape of Australopithecus sediba suggests a single variable species.

The fossils from Malapa cave, South Africa, attributed to Australopithecus sediba, include two partial skeletons-MH1, a subadult, and MH2, an adult. Previous research noted differences in the mandibular rami of these individuals. This study tests three hypotheses that could explain these differences. The first two state that the differences are due to ontogenetic variation and sexual dimorphism, respectively. The third hypothesis, which is relevant to arguments suggesting that MH1 belongs in the genus Australopithecus and MH2 in Homo, is that the differences are due to the two individuals representing more than one taxon. To test these hypotheses, we digitized two-dimensional sliding semilandmarks in samples of Gorilla, Pan, Pongo, and Homo, as well as MH1 and MH2. We document large amounts of shape variation within all extant species, which is related neither to ontogeny nor sexual dimorphism. Extant species nevertheless form clusters in shape space, albeit with some overlap. The shape differences in extant taxa between individuals in the relevant age categories are minimal, indicating that it is unlikely that ontogeny explains the differences between MH1 and MH2. Similarly, the pattern of differences between MH1 and MH2 is inconsistent with those found between males and females in the extant sample, suggesting that it is unlikely that sexual dimorphism explains these differences. While the difference between MH1 and MH2 is large relative to within-species comparisons, it does not generally fall outside of the confidence intervals for extant intraspecific variation. However, the MH1-MH2 distance also does not plot outside and below the between-species confidence intervals. Based on these results, as well as the contextual and depositional evidence, we conclude that MH1 and MH2 represent a single species and that the relatively large degree of variation in this species is due to neither ontogeny nor sexual dimorphism.

Is Central Asia the eastern outpost of the Neandertal range? A reassessment of the Teshik-Tash child.

Since its discovery in southeastern Uzbekistan in 1938, the Teshik-Tash child has been considered a Neandertal. Its affinity is important to studies of Late Pleistocene hominin growth and development as well as interpretations of the Central Asian Middle Paleolithic and the geographic distribution of Neandertals. A close examination of the original Russian monograph reveals the incompleteness of key morphologies associated with the cranial base and face and problems with the reconstruction of the Teshik-Tash cranium, making its Neandertal attribution less certain than previously assumed. This study reassesses the Neandertal status of Teshik-Tash 1 by comparing it to a sample of Neandertal, Middle and Upper Paleolithic modern humans, and recent human sub-adults. Separate examinations of the cranium and mandible are conducted using multinomial logistic regression and discriminant function analysis to assess group membership. Results of the cranial analysis group Teshik-Tash with Upper Paleolithic modern humans when variables are not size-standardized, while results of the mandibular analysis place the specimen with recent modern humans for both raw and size-standardized data. Although these results are influenced by limitations related to the incomplete nature of the comparative sample, they suggest that the morphology of Teshik-Tash 1 as expressed in craniometrics is equivocal. Although, further quantitative studies as well as additional sub-adult fossil finds from this region are needed to ascertain the morphological pattern of this specimen specifically, and Central Asian Middle Paleolithic hominins in general, these results challenge current characterizations of this territory as the eastern boundary of the Neandertal range during the Late Pleistocene.

A fine line: a comparison of methods for estimating ages of linear enamel hypoplasia formation.

It is accepted that linear enamel hypoplasias (LEHs), a specific type of enamel thickness deficiency, are related to periodic physiological disruptions to enamel matrix secretion during times that teeth are developing. Thus, LEHs are treated as general indicators of metabolic stress. Because the disruptions that cause LEHs affect only the portion of the crown that is in the process of forming, determining their locations allows researchers to reconstruct chronologies of stressful events. It is widely held that the many of the commonly used macroscopic methods for estimating the timing of LEHs are imprecise and do not conform to our current understanding of the process of enamel formation. The goal of the present study is to compare estimated ages of LEH formation produced by two of the most commonly used macroscopic methods to those derived from data in recent histological studies that include more precise information about the timing of crown formation across diverse human populations. These approaches are compared in two ways: 1) by creating a theoretical model using simulated LEHs and 2) empirically, by analyzing data collected on a sample of ancient Nubians from Semna South (present-day Sudan). Results indicate that the approach derived from histological studies provides significantly higher age estimates than the commonly used methods and this difference is particularly marked in early forming LEHs. The magnitude of this difference is large enough to produce divergent interpretation of bioarchaeological datasets and suggests that reevaluation of the methods used to estimate ages of LEH formation may be justified.

Ontogenetic variation in the mandibular ramus of great apes and humans.

Considerable variation exists in mandibular ramus form among primates, particularly great apes and humans. Recent analyses of adult ramal morphology have suggested that features on the ramus, especially the coronoid process and sigmoid notch, can be treated as phylogenetic characters that can be used to reconstruct relationships among great ape and fossil hominin taxa. Others have contended that ramal morphology is more influenced by function than phylogeny. In addition, it remains unclear how ontogeny of the ramus contributes to adult variation in great apes and humans. Specifically, it is unclear whether differences among adults appear early and are maintained throughout ontogeny, or if these differences appear, or are enhanced, during later development. To address these questions, the present study examined a broad ontogenetic sample of great apes and humans using two-dimensional geometric morphometric analysis. Variation within and among species was summarized using principal component and thin plate spline analyses, and Procrustes distances and discriminant function analyses were used to statistically compare species and age classes. Results suggest that morphological differences among species in ramal morphology appear early in ontogeny and persist into adulthood. Morphological differences among adults are particularly pronounced in the height and angulation of the coronoid process, the depth and anteroposterior length of the sigmoid notch, and the inclination of the ramus. In all taxa, the ascending ramus of the youngest specimens is more posteriorly inclined in relation to the occlusal plane, shifting to become more upright in adults. These results suggest that, although there are likely functional influences over the form of the coronoid process and ramus, the morphology of this region can be profitably used to differentiate among great apes, modern humans, and fossil hominid taxa.

New hominin remains from Uzbekistan.

Although the Paleolithic occupations of Uzbekistan and the neighboring foothill regions of Tajikistan and Kazakhstan are well-documented, almost no hominin fossil material has been discovered in the area since Teshik-Tash 1 in 1938. Here we describe and offer a preliminary comparative framework for hominin remains that were recovered in 2003 from two Middle Paleolithic sites in Uzbekistan, Obi-Rakhmat Grotto and Anghilak Cave. The description of Teshik-Tash as a Neandertal and the preponderance of lithic assemblages identified as Mousterian in character has supported the interpretation of the region as the eastern-most extent of the Neandertal range. The material from Obi-Rakhmat (OR-1), a subadult represented by part of a permanent maxillary dentition and a fragmentary cranium, expresses a relatively Neandertal-like dentition coupled with more ambiguous cranial anatomy. The remains from Anghilak Cave include a non-diagnostic, diminutive right fifth metatarsal (AH-1). These findings are important additions to the Central Asia hominin fossil record.