Further analyses of the structural organization of Homo luzonensis teeth: Evolutionary implications.

The species Homo luzonensis has recently been described based on a set of dental and postcranial elements found at Callao Cave (Northern Luzon, Philippines) and dated to at least 50-67 ka. Seven postcanine maxillary teeth are attributed to this taxon, five of them belonging to the same individual (CCH6) and representing the holotype of H. luzonensis, whereas the isolated upper premolar CCH8 and the upper third molar CCH9 are paratypes of the species. The teeth are characterized by their small dimensions associated with primitive features, as also found in Homo floresiensis, another hominin having evolved in an insular environment of Southeast Asia. Postcranial bones of the hands and feet of H. luzonensis and H. floresiensis show Homo habilis-like or australopith-like features, whereas cranial and dental morphology are more consistent with the Asian Homo erectus morphology. Due to this mosaic morphology, the origin and phylogenetic relationships of both H. luzonensis and H. floresiensis are still debated. To test the hypotheses that H. luzonensis derives from H. erectus or from an earlier small-brained hominin, we analyzed the µCT scans of the teeth. We investigated both external and internal tooth structure using morphometric methods including: crown outline shape, tooth crown tissue proportions, enamel-dentine junction shape, and pulp morphology. Homo luzonensis external crown morphology aligns more with H. erectus than with H. habilis/H. rudolfensis. The internal structural organization of H. luzonensis teeth exhibits more affinities with that of H. erectus and H. floresiensis than with Neanderthals and modern humans. Our results suggest that both H. floresiensis and H. luzonensis likely evolved from some H. erectus groups that dispersed in the various islands of this region and became isolated until endemic speciation events occurred at least twice during the Pleistocene in insular environments.

The first evidence for Late Pleistocene dogs in Italy.

The identification of the earliest dogs is challenging because of the absence and/or mosaic pattern of morphological diagnostic features in the initial phases of the domestication process. Furthermore, the natural occurrence of some of these characters in Late Pleistocene wolf populations and the time it took from the onset of traits related to domestication to their prevalence remain indefinite. For these reasons, the spatiotemporal context of the early domestication of dogs is hotly debated. Our combined molecular and morphological analyses of fossil canid remains from the sites of Grotta Paglicci and Grotta Romanelli, in southern Italy, attest of the presence of dogs at least 14,000 calibrated years before present. This unambiguously documents one of the earliest occurrence of domesticates in the Upper Palaeolithic of Europe and in the Mediterranean. The genetic affinity between the Palaeolithic dogs from southern Italy and contemporaneous ones found in Germany also suggest that these animals were an important common adjunct during the Late Glacial, when strong cultural diversification occurred between the Mediterranean world and European areas north of the Alps. Additionally, aDNA analyses indicate that this Upper Palaeolithic dog lineage from Italy may have contributed to the genetic diversity of living dogs.

A new species of Homo from the Late Pleistocene of the Philippines.

A hominin third metatarsal discovered in 2007 in Callao Cave (Northern Luzon, the Philippines) and dated to 67 thousand years ago provided the earliest direct evidence of a human presence in the Philippines. Analysis of this foot bone suggested that it belonged to the genus Homo, but to which species was unclear. Here we report the discovery of twelve additional hominin elements that represent at least three individuals that were found in the same stratigraphic layer of Callao Cave as the previously discovered metatarsal. These specimens display a combination of primitive and derived morphological features that is different from the combination of features found in other species in the genus Homo (including Homo floresiensis and Homo sapiens) and warrants their attribution to a new species, which we name Homo luzonensis. The presence of another and previously unknown hominin species east of the Wallace Line during the Late Pleistocene epoch underscores the importance of island Southeast Asia in the evolution of the genus Homo.

Maturation of the human foetal basioccipital: quantifying shape changes in second and third trimesters using elliptic Fourier analysis.

During prenatal development, the brain is considered the best maturation criterion for the estimation of foetal physiological age, regardless of the conditions of pregnancy. Unfortunately, the brain lyses very quickly after death, but fortunately, the brain also has a major influence over osseous structures of the cranial base during development. Therefore, we considered the osseous structures of the cranial base potential indirect maturation indicators of foetal age. Because of its early formation and robustness, the basioccipital is a cranial base bone that is often used for studies in biological anthropology. Studies generally use conventional morphometry and bone size ratio to highlight morphological changes occurring during the foetal period and to create age estimation methods. These methods usually define thresholds beyond which the morphology of the basioccipital changes, but do not fully consider the form that might be valuable precisely to visualize its development or improve age estimation methods. Using geometric morphometric methods, the present study aims to analyse the development of the basioccipital during the second and third trimesters of foetal life by quantifying and visualizing shape changes in the inferior view. Basioccipital shapes are used as direct indicators of the maturation of the cranial base and as indirect indicators of the maturation of the brain and, by extension, the whole body. A sample of 221 anonymized computed tomographic (CT) scans of normal foetuses, ranging from 18 to 41 gestational weeks (GW), was used. Elliptic Fourier analysis (EFA) was used to quantify the basioccipital outline, and maturation stages were established to visualize shape changes with a principal component analysis. Our study allowed us precisely to quantify and continuously visualize shape changes occurring during prenatal life. Additionally, this study provides the first evidence of two distinct linear shape trajectories of the basioccipital. Foetuses aged between 18 and 26 GW have a rapid shape change with well-individualized stages, whereas shape changes are less visible in the second trajectory (27-41 GW). Furthermore, intra-stage shape variation is higher for the basioccipital at the beginning of the second and third trimesters than at the first trimester. By using geometric morphometric methods and EFA, this study shows that it was possible to go beyond classical methods. Indeed, the developed methodology enabled the first quantification of the overall shape changes of the basioccipital between gestational ages. The morphological shape changes throughout the foetal period can be useful for anthropological studies and provide new perspectives for immature age estimation methods.

Dental phenotypic shape variation supports a multiple dispersal model for anatomically modern humans in Southeast Asia.

The population history of anatomically modern humans (AMH) in Southeast Asia (SEA) is a highly debated topic. The impact of sea level variations related to the Last Glacial Maximum (LGM) and the Neolithic diffusion on past population dispersals are two key issues. We have investigated competing AMH dispersal hypotheses in SEA through the analysis of dental phenotype shape variation on the basis of very large archaeological samples employing two complementary approaches. We first explored the structure of between- and within-group shape variation of permanent human molar crowns. Second, we undertook a direct test of competing hypotheses through a modeling approach. Our results identify a significant LGM-mediated AMH expansion and a strong biological impact of the spread of Neolithic farmers into SEA during the Holocene. The present work thus favors a "multiple AMH dispersal" hypothesis for the population history of SEA, reconciling phenotypic and recent genomic data.

Analysis of Hyoid-Larynx Complex Using 3D Geometric Morphometrics.

The aim of this study was to obtain a quantitative anatomical description of the hyoid bone-larynx complex using modern 3D reconstruction tools. The study was conducted on 104 bones from CT scan images of living adult subjects. Three-dimensional reconstructions were created from CT scan images using AVIZO 6.2 software package. A study of this complex was carried out using metric and morphological analyses. Characteristics of the hyoid bone and larynx were highly heterogeneous and were closely linked with the sex, height, and weight of the individuals. Height and width of larynx were significantly greater in men than in women (24.99 vs. 17.3 mm, p ≤ 0.05 and 46.75 vs. 41.07, p ≤ 0.05), whereas the thyroid angle was larger in females (81.12° vs. 74.48°, p ≤ 0.05). There was a significant correlation between the height and weight of subjects and different measurements of the hyoid-larynx complex. (Pearson's coefficient correlation r = 0.42, p ≤ 0.05 between the height of thyroid ala and the height of subjects and r = 0.1, p ≤ 0.05 between the height of thyroid ala and the weight of subjects). Shape and size analysis of the hyoid-larynx complex showed the existence of a significant sexual dimorphism and high interindividual heterogeneity depending to patient morphology. These results encourage us to go further with functional and imaging correlations.

Technical Note: Anatomic identification of isolated modern human molars: testing Procrustes aligned outlines as a standardization procedure for elliptic fourier analysis.

The determination of the precise position of permanent first and second modern human molars, following standard tooth identification criteria, is often difficult because of their morphological similarities. Here, we proposed to evaluate the suitability of two-dimensional crown contour shape analysis in achieving this objective. The method was tested separately on 180 first and second maxillary molars (UM) and 180 first and second mandibular molars (LM) securely identified (in anatomical position in their sockets). Generalized Procrustes superimposition is used to normalize the outlines prior to applying elliptic Fourier analyses ("EFAproc" method). Reliability and effectiveness of this morphometric procedure was evaluated by comparing the results obtained for the same dataset with four other morphometric methods of contour analysis. Cross-validated ("leave one individual out") percentages of misclassification yielded by linear discriminant analyses were used for determining the anatomic position of modern human molars. The percentages of misclassifications obtained from every method of contour analysis were low (1.67% to 3.33% for the UM, 5.56% to 6.67% for the LM) indicating the high suitability of crown contour analyses in correctly identifying molars. A reliable protocol, based on predictive linear discriminant analyses, was then proposed for identification of isolated molars. In addition, our results confirmed that the EFAproc method is suitable for normalizing outlines prior to undertaking elliptic Fourier analyses, especially in the case of nearly circular outlines: it obtained better classification than the classic method of normalization of Fourier descriptors for UM and provided also some advantages over the three landmarks-based methods tested here.

"Small size" in the Philippine human fossil record: is it meaningful for a better understanding of the evolutionary history of the negritos?

"Pygmy populations" are recognized in several places over the world, especially in Western Africa and in Southeast Asia (Philippine "negritos," for instance). Broadly defined as "small-bodied Homo sapiens" (compared with neighboring populations), their origins and the nature of the processes involved in the maintenance of their phenotype over time are highly debated. Major results have been recently obtained from population genetics on present-day negrito populations, but their evolutionary history remains largely unresolved. We present and discuss the Upper Pleistocene human remains recovered from Tabon Cave and Callao Cave in the Philippines, which are potentially highly relevant to these research questions. Human fossils have been recovered in large numbers from Tabon Cave (Palawan Island) but mainly from reworked and mixed sediments from several archaeological layers. We review and synthesize the long and meticulous collaborative work done on the archives left from the 1960s excavations and on the field. The results demonstrate the long history of human occupations in the cave, since at least ~30,000 BP. The examination of the Tabon human remains shows a large variability: large and robust for one part of the sample, and small and gracile for the other part. The latter would fit quite comfortably within the range of variation of Philippine negritos. Farther north, on Luzon Island, the human third metatarsal recently recovered from Callao Cave and dated to ~66,000 BP is now the oldest direct evidence of human presence in the Philippines. Previous data show that, compared with H. sapiens (including Philippine negritos), this bone presents a very small size and several unusual morphological characteristics. We present a new analytical approach using three-dimensional geometric morphometrics for comparing the Callao fossil to a wide array of extant Asian mammals, including nonhuman primates and H. sapiens. The results demonstrate that the shape of the Callao metatarsal is definitely closer to humans than to any other groups. The fossil clearly belongs to the genus Homo; however, it remains at the margin of the variation range of H. sapiens. Because of its great antiquity and the presence of another diminutive species of the genus Homo in the Wallace area during this time period (H. floresiensis), we discuss here in detail the affinities and potential relatedness of the Callao fossil with negritos that are found today on Luzon Island.