Florent Détroit

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 Wallaces 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.

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

Significance Current consensus indicates that modern humans originated from an ancestral African population between ∼100–200 ka. The ensuing dispersal pattern is controversial, yet has important implications for the demographic history and genetic/phenotypic structure of extant human populations. We test for the first time to our knowledge the spatiotemporal dimensions of competing out-of-Africa dispersal models, analyzing in parallel genomic and craniometric data. Our results support an initial dispersal into Asia by a southern route beginning as early as ∼130 ka and a later dispersal into northern Eurasia by ∼50 ka. Our findings indicate that African Pleistocene population structure may account for observed plesiomorphic genetic/phenotypic patterns in extant Australians and Melanesians. They point to an earlier out-of-Africa dispersal than previously hypothesized. 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.

“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.

Notes on the Morphology and Age of the Tabon Cave Fossil Homo sapiens

eusebio dizon, florent détroit , françois s émah, christophe falguères , s ébastien hameau, wilfredo ronquillo, and emmanuel cabanis Archaeology Division, National Museum of the Philippines, Manila, The Philippines (Dizon, Ronquillo)/Laboratoire de Préhistoire du Muséum national d’Histoire naturelle et Laboratoire d’Anthropologie de l’Université de la Méditerranée, UMR 6569 du CNRS, IPH, 1, rue René Panhard, F75013 Paris, France (fdetroit@cimrs1.mnhn.fr) (Détroit, Sémah, Falguères, Hameau)/Service de Radiologie de l’Hôpital des Quinze-Vingt et UMR 6569 du CNRS, IPH, 1, rue René Panhard, F-75013 Paris, France (Cabanis). 14 i 02

Small Size in the Philippine Human Fossil Record: Is It Meaningful for a Better Understanding of the Evolutionary History of the Negritos?

Abstract “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.

First description of the Cro-Magnon 1 endocast and study of brain variation and evolution in anatomically modern Homo sapiens

Paleoneurology is an important research field for studies of human evolution. Variations in the size and shape of the endocranium are a useful means of distinguishing between different hominin species, while brain asymmetry is related to behaviour and cognitive capacities. The evolution of the hominin brain is well documented and substantial literature has been produced on this topic, mostly from studies of endocranial casts, or endocasts. However, we have only little information about variations in endocranial form, size and shape in fossil anatomically modern Homo sapiens (AMH) and about the evolution of the brain since the emergence of our species. One good illustration of this limited knowledge is that one of the first fossil H. sapiens discovered, in 1868, that is also one of the oldest well-preserved European specimen has never been studied in what concerns its endocranial morphology. The first aim of this study was to propose a detailed description of the endocranial anatomy of Cro-Magnon 1, using imaging methodologies, including an original methodology to quantify endocranial asymmetries. The second aim was to compare samples of the fossil and extant AMH in order to document differences in the form, size and shape of the endocasts. A decrease in absolute endocranial size since the Upper Palaeolithic was noticeable. Although both extant and older endocrania have the same anatomical layout, we nonetheless found non-allometric differences in the relative size and organization of different parts of the brain. These document previously unknown intraspecific anatomical variations in the H. sapiens brain, demonstrating its plasticity, with some areas (frontal and occipital lobes) having been more subject to variation than others (parietal, temporal or cerebellar lobes). That may be due to constraints to maintain an optimal performance while reducing in size and changing in shape during our recent evolution.RésuméLa paléoneurologie est un champ de recherche important dans le cadre des études sur l’évolution humaine. Les variations de taille et de forme de l’endocrâne sont en effet utiles pour différencier les différentes espèces d’homininés, alors que les asymétries cérébrales sont reliées au comportement et aux capacités cognitives. Pourtant, notre connaissance de l’évolution et de la variation du cerveau d’Homo sapiens, depuis l’apparition de notre espèce, est très lacunaire. Dans un premier temps, nous détaillons l’anatomie et les asymétries (en proposant une méthode innovante de quantification de ces dernières) de l’endocrâne de Cro-Magnon 1, un des représentants européens les mieux conservés et les plus anciens des Hommes anatomiquement modernes, qui n’avait encore pu être analysé. Puis, une étude comparative entre un échantillon de spécimens fossiles et actuels d’Homo sapiens est effectuée. Bien qu’un substrat anatomique commun soit présent, certaines différences de taille et d’organisation ont été observées entre ces deux échantillons. Ces résultats illustrent la plasticité du cerveau au sein de notre espèce et documentent une variabilité anatomique encore inconnue.

Première description de l’endocrâne de Cro-Magnon 1 et étude de la variation et de l’évolution du cerveau chez les Hommes anatomiquement modernes

Paleoneurology is an important research field for studies of human evolution. Variations in the size and shape of the endocranium are a useful means of distinguishing between different hominin species, while brain asymmetry is related to behaviour and cognitive capacities. The evolution of the hominin brain is well documented and substantial literature has been produced on this topic, mostly from studies of endocranial casts, or endocasts. However, we have only little information about variations in endocranial form, size and shape in fossil anatomically modern Homo sapiens (AMH) and about the evolution of the brain since the emergence of our species. One good illustration of this limited knowledge is that one of the first fossil H. sapiens discovered, in 1868, that is also one of the oldest well-preserved European specimen has never been studied in what concerns its endocranial morphology. The first aim of this study was to propose a detailed description of the endocranial anatomy of Cro-Magnon 1, using imaging methodologies, including an original methodology to quantify endocranial asymmetries. The second aim was to compare samples of the fossil and extant AMH in order to document differences in the form, size and shape of the endocasts. A decrease in absolute endocranial size since the Upper Palaeolithic was noticeable. Although both extant and older endocrania have the same anatomical layout, we nonetheless found non-allometric differences in the relative size and organization of different parts of the brain. These document previously unknown intraspecific anatomical variations in the H. sapiens brain, demonstrating its plasticity, with some areas (frontal and occipital lobes) having been more subject to variation than others (parietal, temporal or cerebellar lobes). That may be due to constraints to maintain an optimal performance while reducing in size and changing in shape during our recent evolution.RésuméLa paléoneurologie est un champ de recherche important dans le cadre des études sur l’évolution humaine. Les variations de taille et de forme de l’endocrâne sont en effet utiles pour différencier les différentes espèces d’homininés, alors que les asymétries cérébrales sont reliées au comportement et aux capacités cognitives. Pourtant, notre connaissance de l’évolution et de la variation du cerveau d’Homo sapiens, depuis l’apparition de notre espèce, est très lacunaire. Dans un premier temps, nous détaillons l’anatomie et les asymétries (en proposant une méthode innovante de quantification de ces dernières) de l’endocrâne de Cro-Magnon 1, un des représentants européens les mieux conservés et les plus anciens des Hommes anatomiquement modernes, qui n’avait encore pu être analysé. Puis, une étude comparative entre un échantillon de spécimens fossiles et actuels d’Homo sapiens est effectuée. Bien qu’un substrat anatomique commun soit présent, certaines différences de taille et d’organisation ont été observées entre ces deux échantillons. Ces résultats illustrent la plasticité du cerveau au sein de notre espèce et documentent une variabilité anatomique encore inconnue.

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

Significance The question of the origin of Pacific Islanders has exercised scientists since the first explorers’ voyages of the 16th century. Physical resemblance between Polynesians and Asian populations was detected, but in between were the islands of Melanesia, inhabited largely by people of different phenotype. However, the Lapita culture bridged this geographical divide 3,000 y ago. Morphological studies of early Lapita colonists from Teouma, Vanuatu align them with present-day Polynesian and Asian populations, whereas skeletal remains of later generations show a more Melanesian phenotype predominating. We suggest that migration streams from already-inhabited parts of Melanesia dating from the late-Lapita phase ultimately dominated the original Polynesian phenotype in eastern Melanesia, but not in Polynesia, which became relatively isolated soon after initial settlement. 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.

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.

Fossil hominins, quadrupedal primates and the origin of human bipedalism: a 3D geometric morphometric analysis of the Primate hamate

This note illustrates the value of studying non-human primates, especially quadrupedal primates, in order to investigate the origins of human bipedalism. Two distinct hypotheses postulate that hominins and African great apes share a common ancestor predominantly engaged in specialized forms of locomotion, i.e., arboreal orthogrady (climbing or arboreal bipedalism) on the one hand and semiterrestrial locomotion (which includes climbing and quadrupedalism) on the other. Both hypotheses are supported by analysis of the wrist morphology of Pliocene hominins, and both have recently been challenged by a third hypothesis based on the study of Ardipithecus ramidus wrist morphology, which has shown general affinities between the latter hominin and quadrupedal primates. However, all three interpretations rely on rather limited knowledge of the variability of wrist bones in quadrupedal primates. Here, we propose to address the question of the origins of human bipedalism by means of a three-dimensional analysis of a carpal bone, the hamate, whose morphology appears to vary according to the locomotor behaviour of primates. We compared the original specimens of Pliocene hominins (Australopithecus) with a large sample of non-human primates, including various quadrupedal anthropoids. Our results confirm that, on the one hand, the shape of the hamate in primates varies significantly according to their locomotor behaviour and, on the other hand, that the hypothesis of the semiterrestrial origin of human bipedalism can be rejected. The affinities between Pliocene hominins and most of extant quadrupedal primates indicate that the hands of early hominins partly retained a morphology inherited from a generalist quadrupedal ancestor, which concurs with the hypothesis recently proposed from the hand bones of Ar. ramidus.RésuméCette note vise à illustrer l’intérêt d’étudier les primates non-humains, notamment quadrupèdes, pour mieux caractériser l’origine de la bipédie humaine. Deux hypothèses stipulent que les homininés partageraient avec les grands singes africains un ancêtre commun impliqué majoritairement dans une forme de locomotion spécialisée, à savoir : l’hypothèse d’une orthogradie arboricole (grimper ou bipédie arboricole) et l’hypothèse d’une semi-terrestrialité (qui inclue quadrupédie et grimper). Ces deux propositions sont notamment supportées par l’analyse morphologique du poignet des homininés pliocènes. Ces propositions ont été récemment remises en cause par une troisième interprétation fondée sur l’étude morphologique des os du poignet d’Ardipithecus ramidus, et qui a mis en évidence des affinités globales entre ce dernier homininé et des singes quadrupèdes. Cependant, ces trois propositions reposent sur une connaissance limitée de la variabilité de ces os chez les primates quadrupèdes. Nous proposons donc d’aborder la question de l’origine de la bipédie humaine par l’analyse tri-dimensionnelle d’un os carpien, l’hamatum, dont la morphologie varierait selon les modes locomoteurs des primates, en considérant des fossiles originaux d’homininés (Australopithecus) ainsi qu’un large échantillon d’anthropoïdes actuels, incluant une grande variété de primates quadrupèdes. Nos résultats confirment d’une part, que la forme de l’hamatum des primates varie selon les comportements locomoteurs, et d’autre part, que l’hypothèse d’une origine semiterrestre de la bipédie humaine peut être rejetée. Les affinités entre les homininés pliocènes et la plupart des primates quadrupèdes actuels soutiennent que les mains des homininés anciens ont en partie retenu une morphologie héritée d’un primate quadrupède généraliste. Ce résultat est en accord avec l’hypothèse récemment proposée à partir des os de la main d’Ar. ramidus.