"Feeling at home in Vanuatu": Integration of newcomers from the East during the last millennium.

Several localities across the Vanuatu archipelago (Melanesia), so-called 'Polynesian Outliers', are inhabited by communities that display Polynesian linguistic and cultural features although being located outside the Polynesian Triangle. Several introductions of Polynesian genetic components to Central and Southern Vanuatu during the last millenium have resulted in the cultural distinctiveness observed among the Polynesian Outliers in Vanuatu. However, social, political or economic process surrounding the exchange of genes between Polynesian and local individuals remain unidentified. Recent bioanthropological studies suggest the existence of female mobilities from neighboring regions to Vanuatu but also to the Polynesian Outliers of Taumako (Solomon Islands) within patrilocal societies. We aim to examine the hypothesis that Polynesian biological affinities observed in ancient individuals from Vanuatu are gendered or sex-specific, and that some of the Polynesian migrations during the last millennium may have involved practices of exogamy. By reconstructing phenotypes and biological identities from 13 archaeologically-recovered human skulls (400-300 years ago) from "Polynesian-related" regions of Vanuatu, we provide new insights to better contextualize the settlement patterns of Polynesian individuals. Eastern-Pacific associated phenotype are observable in 4 women from the Eretok burial complex (Efate region) and the Polynesian Outlier of Futuna, who were buried in close proximity to individuals with Western-Pacific associated phenotype. We suggest that close integration of individuals from the East into the local Vanuatu society, as well as the practice of exogamy, might have been key processes contributing to the preservation of Polynesian cultural features in Vanuatu over the past millennium. Our finding are cross-referenced with oral records from these two areas, as well as the known genetic makeup of the Vanuatu Polynesian Outliers.

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.

Covariation of proximal finger and toe phalanges in Homo sapiens: A novel approach to assess covariation of serially corresponding structures.

OBJECTIVES: As hands and feet are serially repeated corresponding structures in tetrapods, the morphology of fingers and toes is expected to covary due to a shared developmental origin. The present study focuses on the covariation of the shape of proximal finger and toe phalanges of adult Homo sapiens to determine whether covariation is different in the first ray relative to the others, as its morphology is also different. MATERIAL AND METHODS: Proximal phalanges of 76 individuals of unknown sex (Muséum national d'Histoire naturelle, Paris, and the Natural History Museum, London) were digitized using a surface scanner. Landmarks were positioned on 3D surface models of the phalanges. Generalized Procrustes analysis and two-block partial least squares (PLS) analyses were conducted. A novel landmark-based geometric morphometric approach focusing on covariation is based on a PCoA of the angles between PLS axes in morphospace. The results can be statistically evaluated. RESULTS: The difference in PCo scores between the first and the other rays indicates that the integration between the thumb and the big toe is different from that between the lateral rays of the hand and foot. DISCUSSION: We speculate that the results are possibly the evolutionary consequence of differential selection pressure on the big toe relative to the other toes related to the rise of bipedalism, which is proposed to have emerged very early in the hominin clade. In contrast, thumb morphology and its precision grip never ceased undergoing changes, suggesting less acute selection pressures related to the evolution of the precision grip.

Palaeoproteomics gives new insight into early southern African pastoralism.

The advent of domestication is a major step that transformed the subsistence strategies of past human societies. In Africa, domestic caprines (sheep and goat) were introduced in the north-eastern part of the continent from the Near East more than 9000 years ago. However, their diffusion southwards was slow. They are thought to have made their first appearance in the southern part of the continent ca. 2000 years ago, at a few Later Stone Age sites, including Leopard Cave (Erongo region, Namibia), which provided the oldest directly dated remains assigned to sheep or goat on the basis of morphology of bones and teeth. However, similarities in morphology, not only between these two domesticated caprine species, but also between them and the small wild antelopes, raised questions about the morphological species attribution of these remains. Additionally, the high fragmentation of the site's osteological remains makes it difficult to achieve species-level taxonomic identification by comparative anatomy. In this paper, we report molecular species identification of the Leopard Cave remains using palaeoproteomics, a method that uses protein markers in bone and tooth collagen to achieve taxonomic identification of archaeological remains. We also report new direct radiocarbon dates. Wild antelope remains from museum collections were used to enrich the available protein record and propose de novo type I collagen sequences. Our results demonstrate that the remains morphologically described as domesticates actually belong to a wild antelope species and that domestic caprines first appeared at Leopard Cave 1500 years later than previously thought. This study illustrates that the use of palaeoproteomics coupled with direct radiocarbon dates is particularly suited to complement classic zooarchaeological studies, in this case concerning the arrival of the first herding practices in arid environments.

The Middle Pleistocene hominin mandible from Payre (Ardèche, France).

Although Neandertals are the best-known fossil hominins, the tempo and evolutionary processes in their lineage are strongly debated. This is in part due to the scarcity of the fossil record, in particular before the marine isotopic stage (MIS) 5. In 2010, a partial hominin mandible was discovered at the Middle Paleolithic site of Payre (France) in a layer that is dated to the end of MIS 8/beginning of MIS 7, a time period for which very few fossils are known in Europe. The Payre 15 mandible retains the complete symphyseal region and right lateral corpus with heavily worn P4, M1, and M2 in situ. Taphonomic modifications in the form of three notches suggest that this individual was chewed by a carnivore. We provide here the first detailed description of this specimen and a comparative analysis that includes morphological features, linear mandibular dimensions, an elliptic Fourier analysis of the symphysis, and a morphometric analysis of the M1 roots (based on segmented CT scan data). Our comparative sample encompasses European Middle and Upper Pleistocene specimens attributed to Homo heidelbergensis and Homo neanderthalensis, Upper Pleistocene Homo sapiens, and Holocene Homo sapiens. The Payre 15 mandible shows a combination of primitive and Neandertal-like features, with a receding symphyseal profile without any element of the mentum osseum, a posterior location of the mental foramen and lateral prominence. Its mandibular body is tall and thick anteriorly. Payre 15 has mesotaurodont M1 roots and a three-rooted M2. By its dimensions and combination of features, Payre 15 aligns better with Middle Pleistocene European hominins than with MIS 6-3 Neandertals. Noteworthy, it falls well within the range of variation of the Sima de los Huesos sample. Our results underscore that the total pattern of Neandertal-derived morphology was not achieved at the beginning of the MIS 7 and suggest a low level of mandibular diachronic changes for the period MIS 11-7.

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.

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.

Morphometric Assessment of Convergent Tool Technology and Function during the Early Middle Palaeolithic: The Case of Payre, France.

There appears to be little doubt as to the existence of an intentional technological resolve to produce convergent tools during the Middle Palaeolithic. However, the use of these pieces as pointed tools is still subject to debate: i.e., handheld tool vs. hafted tool. Present-day technological analysis has begun to apply new methodologies in order to quantify shape variability and to decipher the role of the morphology of these pieces in relation to function; for instance, geometric morphometric analyses have recently been applied with successful results. This paper presents a study of this type of analysis on 37 convergent tools from level Ga of Payre site (France), dated to MIS 8-7. These pieces are non-standardized knapping products produced by discoidal and orthogonal core technologies. Moreover, macro-wear studies attest to various activities on diverse materials with no evidence of hafting or projectile use. The aim of this paper is to test the geometric morphometric approach on non-standardized artefacts applying the Elliptical Fourier analysis (EFA) to 3D contours and to assess the potential relationship between size and shape, technology and function. This study is innovative in that it is the first time that this method, considered to be a valuable complement for describing technological and functional attributes, is applied to 3D contours of lithic products. Our results show that this methodology ensures a very good degree of accuracy in describing shape variations of the sharp edges of technologically non-standardized convergent tools. EFA on 3D contours indicates variations in deviations of the outline along the third dimension (i.e., dorso-ventrally) and yields quantitative and insightful information on the actual shape variations of tools. Several statistically significant relationships are found between shape variation and use-wear attributes, though the results emphasize the large variability of the shape of the convergent tools, which, in general, does not show a strong direct association with technological features and function. This is in good agreement with the technological context of this chronological period, characterized by a wide diversity of non-standardized tools adapted to multipurpose functions for varied subsistence activities.

Early Lapita skeletons from Vanuatu show Polynesian craniofacial shape: Implications for Remote Oceanic settlement and Lapita origins.

With a cultural and linguistic origin in Island Southeast Asia the Lapita expansion is thought to have led ultimately to the Polynesian settlement of the east Polynesian region after a time of mixing/integration in north Melanesia and a nearly 2,000-y pause in West Polynesia. One of the major achievements of recent Lapita research in Vanuatu has been the discovery of the oldest cemetery found so far in the Pacific at Teouma on the south coast of Efate Island, opening up new prospects for the biological definition of the early settlers of the archipelago and of Remote Oceania in general. Using craniometric evidence from the skeletons in conjunction with archaeological data, we discuss here four debated issues: the Lapita-Asian connection, the degree of admixture, the Lapita-Polynesian connection, and the question of secondary population movement into Remote Oceania.

Genomic and cranial phenotype data support multiple modern human dispersals from Africa and a southern route into Asia.

Despite broad consensus on Africa as the main place of origin for anatomically modern humans, their dispersal pattern out of the continent continues to be intensely debated. In extant human populations, the observation of decreasing genetic and phenotypic diversity at increasing distances from sub-Saharan Africa has been interpreted as evidence for a single dispersal, accompanied by a series of founder effects. In such a scenario, modern human genetic and phenotypic variation was primarily generated through successive population bottlenecks and drift during a rapid worldwide expansion out of Africa in the Late Pleistocene. However, recent genetic studies, as well as accumulating archaeological and paleoanthropological evidence, challenge this parsimonious model. They suggest instead a "southern route" dispersal into Asia as early as the late Middle Pleistocene, followed by a separate dispersal into northern Eurasia. Here we test these competing out-of-Africa scenarios by modeling hypothetical geographical migration routes and assessing their correlation with neutral population differentiation, as measured by genetic polymorphisms and cranial shape variables of modern human populations from Africa and Asia. We show that both lines of evidence support a multiple-dispersals model in which Australo-Melanesian populations are relatively isolated descendants of an early dispersal, whereas other Asian populations are descended from, or highly admixed with, members of a subsequent migration event.

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.

Comparative morphological and morphometric description of the hominin calvaria from Bukuran (Sangiran, Central Java, Indonesia).

We describe the hominin skull (called here "Bukuran") discovered in the lower Kabuh (or "Bapang") series near Sendangbusik, from the Bukuran area in the Sangiran dome. The fossil, heavily mineralized, consists of the parieto-occipital and the left temporal, and the frontal bones. When combined, those two cranial parts represent a rather complete and well-preserved calvaria. Its stratigraphic position was established after the discovery. A detailed description is presented of the morphological and metric features of the Bukuran calvaria, and comparisons are made with Asian Homo erectus from Indonesia and China. The estimated cranial capacity of Bukuran, the general shape of its cranial vault, its ectocranial structures, and its morphological and metrical characters are in the range of Asian Homo erectus, and show clear affinities with other Indonesian members of the species. We discuss the evolutionary status of the Bukuran calvaria and its implication for hominin history on Java.

"Of sheep and men": earliest direct evidence of caprine domestication in southern Africa at Leopard Cave (Erongo, Namibia).

The origins of herding practices in southern Africa remain controversial. The first appearance of domesticated caprines in the subcontinent is thought to be c. 2000 years BP; however, the origin of this cultural development is still widely debated. Recent genetic analyses support the long-standing hypothesis of herder migration from the north, while other researchers have argued for a cultural diffusion hypothesis where the spread of herding practices took place without necessarily implicating simultaneous and large population movements. Here we document the Later Stone Age (LSA) site of Leopard Cave (Erongo, Namibia), which contains confirmed caprine remains, from which we infer that domesticates were present in the southern African region as early as the end of the first millennium BC. These remains predate the first evidence of domesticates previously recorded for the subcontinent. This discovery sheds new light on the emergence of herding practices in southern Africa, and also on the possible southward routes used by caprines along the western Atlantic coast.

New evidence for a 67,000-year-old human presence at Callao Cave, Luzon, Philippines.

Documentation of early human migrations through Island Southeast Asia and Wallacea en route to Australia has always been problematic due to a lack of well-dated human skeletal remains. The best known modern humans are from Niah Cave in Borneo (40-42ka), and from Tabon Cave on the island of Palawan, southwest Philippines (47+/-11ka). The discovery of Homo floresiensis on the island of Flores in eastern Indonesia has also highlighted the possibilities of identifying new hominin species on islands in the region. Here, we report the discovery of a human third metatarsal from Callao Cave in northern Luzon. Direct dating of the specimen using U-series ablation has provided a minimum age estimate of 66.7+/-1ka, making it the oldest known human fossil in the Philippines. Its morphological features, as well as size and shape characteristics, indicate that the Callao metatarsal definitely belongs to the genus Homo. Morphometric analysis of the Callao metatarsal indicates that it has a gracile structure, close to that observed in other small-bodied Homo sapiens. Interestingly, the Callao metatarsal also falls within the morphological and size ranges of Homo habilis and H. floresiensis. Identifying whether the metatarsal represents the earliest record of H. sapiens so far recorded anywhere east of Wallace's Line requires further archaeological research, but its presence on the isolated island of Luzon over 65,000 years ago further demonstrates the abilities of humans to make open ocean crossings in the Late Pleistocene.