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The abundant femoral assemblage of Homo naledi found in the Dinaledi Chamber provides a unique opportunity to test hypotheses regarding the taxonomy, locomotion, and loading patterns of this species. Here we describe neck and shaft cross-sectional structure of all the femoral fossils recovered in the Dinaledi Chamber and compare them to a broad sample of fossil hominins, recent humans, and extant apes. Cross-sectional geometric (CSG) properties from the femoral neck (base of neck and midneck) and diaphysis (subtrochanteric region and midshaft) were obtained through CT scans for H. naledi and through CT scans or from the literature for the comparative sample. The comparison of CSG properties of H. naledi and the comparative samples shows that H. naledi femoral neck is quite derived with low superoinferior cortical thickness ratio and high relative cortical area. The neck appears superoinferiorly elongated because of two bony pilasters on its superior surface. Homo naledi femoral shaft shows a relatively thick cortex compared to the other hominins. The subtrochanteric region of the diaphysis is mediolaterally elongated resembling early hominins while the midshaft is anteroposteriorly elongated, indicating high mobility levels. In term of diaphyseal robusticity, the H. naledi femur is more gracile that other hominins and most apes. Homo naledi shows a unique combination of characteristics in its femur that undoubtedly indicate a species committed to terrestrial bipedalism but with a unique loading pattern of the femur possibly consequence of the unique postcranial anatomy of the species.

The morphology and size of the Neandertal thorax is a subject of growing interest due to its link to general aspects of body size and shape, including physiological aspects related to bioenergetics and activity budgets. However, the number of well-preserved adult Neandertal costal remains is still low. The recent finding of new additional costal remains from the Regourdou 1 (R1) skeleton has rendered this skeleton as one of the most complete Neandertal costal skeletons with a minimum of 18 ribs represented, five of which are complete or virtually complete. Here we describe for the first time all the rib remains from R1 and compare them to a large modern Euroamerican male sample as well as to other published Neandertal individuals. The costal skeleton of this individual shows significant metric and morphological differences from our modern human male comparative sample. The perceived differences include: dorsoventrally large 1st and 2nd ribs, 3rd ribs with a very closed dorsal curvature and large maximum diameters at the posterior angle, a large tubercle-iliocostal line distance in the 4th rib, thick shafts at the dorsal end of its 6th ribs, thick mid-shafts of the 8th ribs, large articular tubercles at the 9th ribs, and thick shafts of the 11th and 12th ribs. Here we also describe a new mesosternal fragment: the left lateral half of sternebral segments 4 and 5. This portion reveals that the mesosternum of R1 had a sternal foramen in its inferiormost preserved sternal segment and supports previous estimation of the total length of this mesosternum. The new costal remains from R1 support the view that Neandertals, when compared with modern humans, show a significantly different thorax, consistent with differences found in other anatomical regions such as the vertebral column and pelvis.

Objectives Humans generally comply with the ecological rule of Allen (1877), with populations from tropical environments exhibiting body proportions in which limb segments are long relative to trunk height compared to temperate groups. This study tests whether ecogeographic differences in intralimb proportions are identifiable among two modern fetal samples of differing ancestry. Materials and methods Data are derived from radiographic measurements of long bone diaphyseal length and crown-heel length (CHL) of contemporary, spontaneously aborted fetuses of African Americans ("black") of assumed African (tropical) ancestry and European Americans ("white") of assumed European (temperate) ancestry (n = 184). Population individual limb elements, brachial, and crural indices are compared via analyses of covariance (ANCOVA). Potential patterns of divergent allometric growth are quantified through principal components analysis (PCA). Results African ancestral distal limb elements were consistently, albeit slightly, longer than those of European ancestry, relative to CHL. None of the ANCOVA interactions with ancestry are statistically significant for limb indices. The radius was the only single element that displayed a statistically significant ancestry effect (p = 0.0435) equating to a 1 mm difference. PCA highlights that upper limbs demonstrate negative allometry and lower limbs demonstrate positive allometry with sample-specific multivariate growth patterns being nearly identical. Differences in growth allometry late in gestation make little contribution to observed differences in adult limb proportions. Discussion No statistically significant ecogeographic patterns were appreciated among intralimb proportions between these groups during the fetal period. This study contributes to a greater appreciation of phenotypic plasticity, ecogeographic variation in ontogeny, and the evolution of modern human diversity.

This paper describes the 108 femoral, patellar, tibial, and fibular elements of a new species of Homo (Homo naledi) discovered in the Dinaledi chamber of the Rising Star cave system in South Africa. Homo naledi possesses a mosaic of primitive, derived, and unique traits functionally indicative of a bipedal hominin adapted for long distance walking and possibly running. Traits shared with australopiths include an anteroposteriorly compressed femoral neck, a mediolaterally compressed tibia, and a relatively circular fibular neck. Traits shared with Homo include a well-marked linea aspera, anteroposteriorly thick patellae, relatively long tibiae, and gracile fibulae with laterally oriented lateral malleoli. Unique features include the presence of two pillars on the superior aspect of the femoral neck and a tubercular distal insertion of the pes anserinus on the tibia. The mosaic morphology of the H. naledi thigh and leg appears most consistent with a species intermediate between Australopithecus spp. and Homo erectus and, accordingly, may offer insight into the nature of the earliest members of genus Homo. These fossils also expand the morphological diversity of the Homo lower limb, perhaps indicative of locomotor diversity in our genus.

International audience; The calcium isotopic composition (δ44/42Ca) of bone and tooth enamel can be used for dietary reconstructions of extant and extinct mammals. In natural conditions, the δ44/42Ca value of bone and teeth varies according to dietary intake with a constant isotopic offset of about −0.6‰. Owing to the poor conservation of collagen, carbon (C), and nitrogen (N) isotopic compositions of the Regourdou Mousterian site (MIS 5, Dordogne, France) previously failed to provide any paleodietary information. Therefore, to reconstruct the trophic chain, we have measured calcium (Ca) isotopes from fossil bone samples of the fauna from the Regourdou site, as well as from three bone samples of the Regourdou 1 Neandertal specimen. The results show a taxon-dependent patterning of the Ca isotopic compositions: herbivores generally have higher δ44/42Ca values than carnivores. All the δ44/42Ca values of Regourdou 1 are low (

In 2015, several disarticulated human skeletal remains were identified in the top layers of the prehistoric site of Buraca da Moira Rock Shelter (Boa Vista, Leiria), during the archaeological excavation carried out under the scope of the EcoPLis — Human Occupations in the Pleistocene Ecotones of the River Lis project. The recovery of chert, constiquartz and quartzite blanks, a schist plate, as well as adornments in bone and shell indicates a Late Neolithic-Chalcolithic chronology. The disarticulated human assemblage, composed of a total of 129 bone and tooth fragments, allowed the estimation of a minimum number of six individuals. Among the remains recovered, an upper portion of an adult right ulna lacking the coronoid process was identified. Replacing it, a semi-oval groovewith smooth contours and exposing some trabecular bone was observed. The location, type of bone change, and the observed signs of bone healing are consistent with an uncommon trauma: a fracture of the coronoid process. In the differential diagnosis, both postmortem changes and developmental disturbances were considered but later excluded. The mechanisms that underlie the bone changes are discussed in light of the clinical and paleopathological literature., No ano de 2015, durante a intervenção arqueológica afeta ao projeto EcoPLis: Ocupação Humana Plistocénica nos Ecótonos do Rio Lis foram identificados vestígios osteológicos humanos desarticulados à superfície e nas camadas iniciais do complexo cársico do Abrigo da Buraca da Moira (Boa Vista, Leiria). Arecuperação de artefactos em quartzo e sílex, de uma placa de xisto, e de adornos em osso e concha sugerem como cronologia o Neolítico Final/Calcolítico. A análise paleobiológica preliminar dos fragmentos ósseos e peças dentárias recuperados (n=129) permitiu inferir que pertencerão a, pelo menos, seis indivíduos. De entre os elementos analisados macroscopicamente, destacou-se um segmento proximal de uma ulna direita pertencente a um indivíduo adulto. Este fragmento caracterizava-se pela ausência do processo coronóide, e pela presença de uma depressão óssea ovalada com contornos suaves, associada a uma ligeira exposição de osso trabecular. A localização, tipo de alteração e a presença de sinais de remodelação óssea são consistentes com uma lesão traumática incomum: a fratura do processo coronóide. No diagnóstico diferencial foram consideradas, e posteriormente excluídas, alterações de cariz tafonómico e problemas de desenvolvimento. Neste trabalho serão discutidos os mecanismos conducentes à lesão à luz da literatura clínica e paleopatológica.

Depuis la fin du XIXe siècle, l’Europe occidentale a fait l’objet de très nombreux travaux pour étudier les modes de subsistance des sociétés préhistoriques. À partir du Paléolithique moyen, les Hommes vont alors s’intéresser au petit gibier, tels que les léporidés, taxons relativement abondants sur ce territoire. Or, distinguer l’origine de leur accumulation dans un site archéologique ne constitue pas une tâche aisée tant les agents pouvant en être responsables sont nombreux (i.e. mortalité naturelle, humains, carnivores terrestres, rapaces nocturnes ou diurnes). Dans cette contribution, nous avons mené des études taphonomique et paléontologique sur les léporidés de Regourdou, gisement moustérien ayant livré le squelette d’un Néandertalien. Les indices recensés suggèrent qu’aucun prédateur n’est intervenu dans les processus d’accumulation des restes de lièvres et de lapins, mais leurs assemblages seraient dus, respectivement, à une mortalité naturelle de type accidentelle et de type attritionnelle. La mise en évidence du premier processus montre que la cavité aurait fonctionné comme un aven-piège à un moment donné de son histoire, tandis que le second révèle l’existence d’une Garenne in situ, interprétée comme la dernière occupation du site durant le MIS 4 ou 3, avant la fin du comblement sédimentaire. Les variations morphologiques et de taille corporelle des individus permettent également de préciser les changements d’ordre paléo-environnementaux depuis le début du fonctionnement du site, confirmant l’attribution des niveaux inférieurs à la période tempérée du MIS 5, et les niveaux supérieurs à une phase plus récente du MIS 4 ou 3. Since the end of the nineteenth century, a great deal of work studying subsistence patterns of prehistoric societies in Western Europe has been done. During the Middle Paleolithic, humans were interested in small game, particularly the Leporidae, taxa that were abundant in their territories. However, distinguishing the exact nature of their origin in an archaeological site is not an easy task, given that numerous agents could be responsible for their accumulation (i.e., natural mortality, acquisition by humans and/or other terrestrial carnivores, or even nocturnal or diurnal raptors). In this contribution, we put forth a new taphonomic and paleontological study of the leporids of Regourdou, a Mousterian site that has yielded a Neandertal skeleton. This study suggests that no predators were involved in the accumulation of the hare or rabbit remains, but rather that they are due to natural (accidental) mortality for the former, and natural (attritional) mortality for the latter. Specifically, the evidence suggests that at one point in its history (for the hares), the cavity functioned as a natural pitfall trap, while for the rabbits it appears to show the existence of a warren, which we interpret as having existed during the last occupation of the site during MIS 4 or 3, before the end of the accumulation of sediments in the site. Morphological variation and body size of individuals also allows us to specify the order of paleoenvironmental changes since the site’s functional beginning, confirming the attribution of the inferior levels to the MIS 5 temperate period, and the upper levels to a more recent phase in MIS 4 or 3.

Aside from the work of Bonifay (see Bonifay et al. 2007 for one of the more recent papers) and various articles following these earlier works (e.g., Binant 1991, Defleur 1993, Maureille et Vandermeersch 2007, Pettitt 2011, see also May 1986 for a more critical analysis), the in situ position of the remains of Regourdou 1 from layer 4 has never actually been discussed on the basis of available data from the salvage operation carried out in October 1957 by E. Bonifay and G. Laplace-Jauretche, under the administrative authority of Francois Bordes, or from the subsequent, more systematic, excavations directed by E. Bonifay between 1961 and 1964. Via the compilation of available information from a number of unpublished documents (Francois Bordes’ field notes, drawings made during the salvage operation, photographs taken in 1957, 1961 and 1962, as well as databases from the 1961 to 1964 excavations), and also a new inventory of human remains (both previously known and recently discovered), it is now possible to more accurately reconstruct the position of the human remains in a Cartesian system. In this, we assume that the concentration of remains uncovered during the salvage operation was in square G2, according to the preliminary systematic excavations carried out in 1961. They also bring to light that while practically no anatomical connections can be demonstrated with any certainty – and despite significant disruptions (all of the hominin remains are spread over 9 squares : G1 to G3, F1 to F3, E1 and E2, D2) – they are mainly positioned in squares G2 and G3 to some degree with respect to the anatomical logic of the human body. We therefore assume that Regourdou 1 was lying flat, with its head to the west – perhaps upon its trunk – close to the wall of the cavity. This result is different from the fetal position hypothesis proposed in Bonifay et al. (2007). Moreover many post-depositional (albeit Pleistocene) disturbances are also evident. We believe that they were likely the result of the utilization and modification of the cavity by brown bears and lagomorphs.Only new excavations at the site, and a better taphonomic understanding of Bonifay’s (1964) layer 4 (in which Regourdou 1 was found), and the exact role of humans in its formation, i.e., their anthropic impact on the layer, will allow us to discuss in more detail the nature of the deposition of the body, and, hopefully, the absence of the skull.

Excepté dans les travaux de E. Bonifay (pour l’un des plus récents, voir Bonifay et al. 2007) et dans différentes contributions qui ont repris ces derniers (p. ex. Binant 1991 ; Defleur 1993 ; Maureille et Vandermeersch 2007 ; Pettitt 2011 voir aussi l’analyse plus critique de May 1986), la position in situ des restes humains de Regourdou 1, provenant de la couche 4 du site n’a jamais été discutée ni sur la base de l’opération de sauvetage réalisée en octobre 1957 (opération dirigée par E. Bonifay et G. Laplace-Jauretche, sous l’autorité administrative de F. Bordes), ni suite aux fouilles programmées dirigées par E. Bonifay entre 1961 et 1964.Après une synthèse des informations disponibles contenues dans de nombreux documents inédits (minutes de terrain de François Bordes, dessins réalisés lors de l’opération de sauvetage, photographies réalisées en 1957 puis en 1961 et 1962, base de données des fouilles 1961-1964) et d’un nouvel inventaire des restes humains (connus et nouvellement découverts), les ossements de Regourdou 1 ont pu être en majorité repositionnés au sein d’un système orthonormé. Ces nouveaux documents permettent de supposer que la concentration de vestiges mis au jour lors de l’opération de sauvetage se situait dans le carré G2 du carroyage des fouilles débutées en 1961.Ils mettent également en évidence que, même si pratiquement aucune connexion anatomique n’est décelée avec certitude et malgré des perturbations très importantes (la totalité des ossements se répartit in fine sur près de neuf carrés : G1 à G3, F1 à F3, E1, E2 et D2), ces restes se distribuent surtout en G2 et en G3 en respectant la logique anatomique du corps humain. Ces observations permettent de suggérer que Regourdou 1 était plutôt en position allongée, la tête à l’ouest - peut-être ramenée sur le tronc - à proximité de la paroi de la cavité. Ce résultat est donc différent de l’hypothèse de la position fœtale proposée dans Bonifay et al. (2007). De plus, de nombreuses perturbations post-dépositionnelles du dépôt initial humain durant le Pléistocène se sont produites probablement en liaison avec la fréquentation de la cavité par l’Ours brun et les lagomorphes.Nous espérons que de nouvelles fouilles du site, et particulièrement l’étude du rôle de l’Homme dans l’accumulation des vestiges de la couche 4 (selon la stratigraphie de Bonifay 1964), nous permettront de discuter des causes de la présence de ce néandertalien et, peut-être, de l’absence de sa boîte crânienne. Aside from the work of Bonifay (see Bonifay et al. 2007 for one of the more recent papers) and various articles following these earlier works (e.g., Binant 1991, Defleur 1993, Maureille et Vandermeersch 2007, Pettitt 2011, see also May 1986 for a more critical analysis), the in situ position of the remains of Regourdou 1 from layer 4 has never actually been discussed on the basis of available data from the salvage operation carried out in October 1957 by E. Bonifay and G. Laplace-Jauretche, under the administrative authority of François Bordes, or from the subsequent, more systematic, excavations directed by E. Bonifay between 1961 and 1964.Via the compilation of available information from a number of unpublished documents (François Bordes’ field notes, drawings made during the salvage operation, photographs taken in 1957, 1961 and 1962, as well as databases from the 1961 to 1964 excavations), and also a new inventory of human remains (both previously known and recently discovered), it is now possible to more accurately reconstruct the position of the human remains in a Cartesian system. In this, we assume that the concentration of remains uncovered during the salvage operation was in square G2, according to the preliminary systematic excavations carried out in 1961. They also bring to light that while practically no anatomical connections can be demonstrated with any certainty – and despite significant disruptions (all of the hominin remains are spread over 9 squares: G1 to G3, F1 to F3, E1 and E2, D2) – are mainly positioned in squares G2 and G3 to some degree with respect to the anatomical logic of the human body. We therefore assume that Regourdou 1 was lying flat, with its head to the west – perhaps upon its trunk – close to the wall of the cavity. This result is different from the fetal position hypothesis proposed in Bonifay et al. (2007). Moreover many post-depositional (albeit Pleistocene) disturbances are also evident. We believe that they were likely the result of the utilization and modification of the cavity by brown bears and lagomorphs.Only new excavations at the site, and a better taphonomic understanding of Bonifay’s (1964) layer 4 (in which Regourdou 1 was found), and the exact role of humans in its formation, i.e., their anthropic impact on the layer, will allow us to discuss in more detail the nature of the deposition of the body, and, hopefully, the absence of the skull.

Dans un article publié dans le n° 20 de la revue Paleo (Madelaine et al. 2008) nous avons annoncé la découverte d’une série de 11 nouveaux restes humains que nous avons pu rapporter à l’individu n° 1 (Vandermeersch et Trinkaus 1995) de Regourdou mis au jour en septembre 1957. Certains, parmi les plus conséquents (fémur, tibia, fibula), représentaient ou complétaient des régions squelettiques supposées manquantes en raison de gestes funéraires (Bonifay et al. 2007).Dans cette contribution, nous augmentons la représentation squelettique de Regourdou 1 avec de nouvelles pièces. Elles proviennent d’une part de la collection de faune (propriété du Musée national de Préhistoire depuis 2002), de celle de la famille Constant et d’autre part, des collections du Musée d’Art et d’Archéologie de Périgueux. Deux ossements, une diaphyse fémorale gauche et un tibia, proviennent des collections de faune du Musée de site de Regourdou. Mais la morphologie du second ne permet pas de le rapporter à l’Homme de Néandertal.Ces nouvelles découvertes nous permettent donc d’enrichir nos connaissances sur la variabilité anatomique néandertalienne et sur l’histoire de l’occupation du site. Elles valident la présence d’au moins un second adulte moustérien, uniquement représenté par un calcaneus droit. Elles nous autorisent aussi à revenir sur une hypothèse intéressant l’histoire taphonomique du sujet le plus complet (Regourdou 1), hypothèse que nous avions avancée en 2008 et qui apparaît désormais fausse. Enfin, l’origine de l’individu représenté par un tibia de morphologie moderne est inconnue. Il nécessitera une tentative de datation absolue directe. In an article published in Paleo number 20 (Madelaine et al. 2008), we announced the discovery of a new series of 11 human remains that we associated with Regourdou individual number 1 (Vandermeersch and Trinkaus 1995), a specimen first brought to light in September 1957. Some of these new bones, in particular the larger ones (femur, tibia, fibula) represent or make more complete skeletal elements that were presumed to have been missing as the result of funerary acts (Bonifay et al. 2007).In this contribution, we increase the skeletal representation of Regourdou 1 with additional new pieces.These come in part from the site’s faunal collections (property of the Musée national de Préhistoire since 2002), but also from the collections of the Musée d’Art et d’Archéologie de Périgueux. In addition, two pieces: a left femoral diaphysis and the proximal half of a tibia come from the collection of the Constant family, but the morphology of the latter does not appear to be that of a Neandertal.These new discoveries increase our knowledge of Neandertal anatomical variability and the history of the site’s occupation, and also permit us to confirm the presence of at least one second adult Mousterian-associated individual, who is thus far represented solely by a right calcaneus. They also allow us to revisit an interesting hypothesis regarding the taphonomic history of the most complete individual from the site (Regourdou 1), a hypothesis we put forth in 2008 that now turns out to be false. Finally, the origin of the individual represented by the tibia (which evinces modern morphology) is unknown. We will therefore need to obtain an absolute direct date on it.

Body mass changes in the Hominini are examined from an evolutionary perspective. Among the conclusions drawn from this examination are the following: 1.) hominids (sensu lato) are among the largest and most terrestrial primates; 2.) early Homo is on average larger in size than Australopithecus (although the two taxa do show some size overlap); 3.) following its emergence in the early Pleistocene, Homo shows a steady increase in body mass, reaching its maximum near the end of the middle Pleistocene; 4.) Homo subsequently decreases in size throughout the late Pleistocene and into the Holocene, only to increase in size again very recently (the so-called “secular trend”); 5.) body mass variation among humans today tends to follow well-known ecogeographical “rules”, which are empirical observations with multiple potential theoretical explanations.

The estimation of human population size during the Pleistocene is complex, and one which has been dealt with extensively in the literature. However, because many of these previous estimations are based in part on archaeological site distributions, they are more a reflection of present-day geography than of what the Earth looked like in the past. We address this issue by calculating an estimation of habitable land area during the Last Glacial Maximum (between 22 and 19 kya) when sea level was 120 m lower than today using the polygon creation function in Google Earth. We then subtract areas of land that were likely uninhabitable during the LGM – either due to glacier cover, extreme aridity, elevation, or areas at high latitudes. From this, the combined habitable land areas of Eurasia, Africa and the Australian landmass are estimated as 76,959,712.4 km 2 . This estimation is then coupled with population density data for medium-to large-bodied carnivores, and ethnographic population density data for hunter–gatherers culled from the literature. Total human census population size in the Old World during the Last Glacial Maximum is estimated at 2,117,000–2,955,000 based on carnivore densities and 3,046,000–8,307,000 for hunter–gatherer densities.

Regourdou is a well-known Middle Paleolithic site which has yielded the fossil remains of a minimum of two Neandertal individuals. The first individual (Regourdou 1) is represented by a partial skeleton while the second one is represented by a calcaneus. The foot remains of Regourdou 1 have been used in a number of comparative studies, but to date a full description and comparison of all the foot remains from the Regourdou 1 Neandertal, coming from the old excavations and from the recent reanalysis of the faunal remains, does not exist. Here, we describe and comparatively assess the Regourdou 1 tarsals, metatarsals and phalanges. They display traits observed in other Neandertal feet, which are different from some traits of the Sima de los Huesos (Atapuerca) hominins and of Middle Paleolithic, Upper Paleolithic and recent modern humans. These Neandertal features are: a rectangular talar trochlea with a large lateral malleolar facet, a broad talar head, a broad calcaneus with a projecting sustentaculum tali, a wide and wedged navicular with a projecting medial tubercle, large and wide bases of the lateral metatarsals, and mediolaterally expanded and robust phalanges that also show hallux valgus in a strongly built hallux.

17 pages; International audience; The understanding of Neanderthal societies, both with regard to their funerary behaviors and their subsistence activities, is hotly debated. Old excavations and a lack of taphonomic context are often factors that limit our ability to address these questions. To better appreciate the exact nature of what is potentially the oldest burial in Western Europe, Regourdou (Montignac-sur-Vézère, Dordogne), and to better understand the taphonomy of this site excavated more than 50 years ago, we report in this contribution a study of the most abundant animals throughout its stratigraphy: the European rabbit (Oryctolagus cuniculus). In addition to questions surrounding the potential bioturbation of the site's stratigraphy, analysis of the Regourdou rabbits could provide new information on Neandertal subsistence behavior. The mortality profile, skeletal-part representation, breakage patterns, surface modification, and comparison with modern reference collections supports the hypothesis that the Regourdou rabbit remains were primarily accumulated due to natural (attritional) mortality. Radiocarbon dates performed directly on the rabbit remains give ages ranging within the second half of Marine Isotope Stage 3, notably younger than the regional Mousterian period. We posit that rabbits dug their burrows within Regourdou's sedimentological filling, likely inhabiting the site after it was filled. The impact of rabbit activity now brings into question both the reliability of the archaeostratigraphy of the site and the paleoenvironmental reconstructions previously proposed for it, and suggests rabbits may have played a role in the distribution of the Neandertal skeletal remains.

Homo erectus and later humans have enlarged body sizes, reduced sexual dimorphism, elongated lower limbs, and increased encephalization compared to Australopithecus , together suggesting a distinct ecological pattern. The mosaic expression of such features in early Homo , including Homo habilis , Homo rudolfensis , and some early H. erectus , suggests that these traits do not constitute an integrated package. We examined the evidence for body mass, stature, limb proportions, body size and dental size dimorphism, and absolute and relative brain size in Homo naledi as represented in the Dinaledi Chamber sample. H. naledi stature and body mass are low compared to reported values for H. erectus , with the exception of some of the smaller bodied Dmanisi H. erectus specimens, and overlap with larger Australopithecus and early Homo estimates . H. naledi endocranial volumes (465–560 cc) and estimates of encephalization quotient are also similar to Australopithecus and low compared to all Homo specimens, with the exception of Homo floresiensis (LB1) and the smallest Dmanisi H. erectus specimen (D4500). Unlike Australopithecus , but similar to derived members of genus Homo , the Dinaledi assemblage of H. naledi exhibits both low levels of body mass and dental size variation, with an estimated body mass index of sexual dimorphism less than 20%, and appears to have an elongated lower limb. Thus, the H. naledi bauplan combines features not typically seen in Homo species (e.g., small brains and bodies) with those characteristic of H. erectus and more recent Homo species (e.g., reduced mass dimorphism, elongated lower limb).

Depuis la fin du XIXe siècle, l’Europe occidentale a fait l’objet de très nombreux travaux pour étudier les modes de subsistance des sociétés préhistoriques. À partir du Paléolithique moyen, les Hommes vont alors s’intéresser au petit gibier, tels que les léporidés, taxons relativement abondants sur ce territoire. Or, distinguer l’origine de leur accumulation dans un site archéologique ne constitue pas une tâche aisée tant les agents pouvant en être responsables sont nombreux (i.e. mortalité naturelle, humains, carnivores terrestres, rapaces nocturnes ou diurnes). Dans cette contribution, nous avons mené des études taphonomique et paléontologique sur les léporidés de Regourdou, gisement moustérien ayant livré le squelette d’un Néandertalien. Les indices recensés suggèrent qu’aucun prédateur n’est intervenu dans les processus d’accumulation des restes de lièvres et de lapins, mais leurs assemblages seraient dus, respectivement, à une mortalité naturelle de type accidentelle et de type attritionnelle. La mise en évidence du premier processus montre que la cavité aurait fonctionné comme un aven-piège à un moment donné de son histoire, tandis que le second révèle l’existence d’une Garenne in situ, interprétée comme la dernière occupation du site durant le MIS 4 ou 3, avant la fin du comblement sédimentaire. Les variations morphologiques et de taille corporelle des individus permettent également de préciser les changements d’ordre paléo-environnementaux depuis le début du fonctionnement du site, confirmant l’attribution des niveaux inférieurs à la période tempérée du MIS 5, et les niveaux supérieurs à une phase plus récente du MIS 4 ou 3. Since the end of the nineteenth century, a great deal of work studying subsistence patterns of prehistoric societies in Western Europe has been done. During the Middle Paleolithic, humans were interested in small game, particularly the Leporidae, taxa that were abundant in their territories. However, distinguishing the exact nature of their origin in an archaeological site is not an easy task, given that numerous agents could be responsible for their accumulation (i.e., natural mortality, acquisition by humans and/or other terrestrial carnivores, or even nocturnal or diurnal raptors). In this contribution, we put forth a new taphonomic and paleontological study of the leporids of Regourdou, a Mousterian site that has yielded a Neandertal skeleton. This study suggests that no predators were involved in the accumulation of the hare or rabbit remains, but rather that they are due to natural (accidental) mortality for the former, and natural (attritional) mortality for the latter. Specifically, the evidence suggests that at one point in its history (for the hares), the cavity functioned as a natural pitfall trap, while for the rabbits it appears to show the existence of a warren, which we interpret as having existed during the last occupation of the site during MIS 4 or 3, before the end of the accumulation of sediments in the site. Morphological variation and body size of individuals also allows us to specify the order of paleoenvironmental changes since the site’s functional beginning, confirming the attribution of the inferior levels to the MIS 5 temperate period, and the upper levels to a more recent phase in MIS 4 or 3.

Homo rudolfensis Samples from Olduvai Gorge, East Lake Turkana, and Lake Malawi were included in this study. The East Lake Turkana fossils available prior to 2010 were examined first-hand, while for the Olduvai and Lake Malawi fossils and KNM-ER 60000, 62000, and 62003 we relied on original observations on fossils and casts as well as published reports (Schrenk et al., 1993; Blumenschine et al., 2003; Leakey et al., 2012). We include the following fossils in the hypodigm of H. rudolfensis: KNM-ER 819, KNM-ER 1470, KNM-ER 1482, KNM-ER 1483, KNM-ER 1590, KNM-ER 1801, KNM-ER 1802, KNM-ER 3732, KNM-ER 3891, KNM-ER 60000, KNM-ER 62000, KNM-ER 62003, OH 65, and UR 501. We do recognize that KNM-ER 60000 and KNM-ER 1802 present some conflicting anatomy that some authors have argued precludes them as conspecific specimens (Leakey et al., 2012); by considering both, we aim to be conservative as they encompass more variation within H. rudolfensis., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Australopithecus africanus The samples attributed to Au. africanus from Taung, Sterkfontein and Makapansgat were employed. Original specimens were examined first-hand by the authors., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Paranthropus boisei Samples from the Omo Shungura sequence, East Lake Turkana, Olduvai Gorge and Konso were included in this study. Original specimens from Olduvai Gorge and East Lake Turkana were examined first-hand, while casts and published reports (Tobias, 1967; Suwa et al., 1996, 1997; Do ḿınguez- Rodrigo et al., 2013) were used to study the Omo and Konso materials. Our postcranial considerations of P. boisei are very limited and we did not rely upon the association of KNM-ER 1500 (Grausz et al., 1988) to derive information about the postcranial skeleton of P. boisei. Paranthropus robustus The samples from Kromdraai, Swartkrans, Sterkfontein, Drimolen, Gondolin, and Coopers were included in this study. First-hand observations of original specimens from all localities were used with the exception of Drimolen fossils, which were compared using published reports (Keyser, 2000; Keyser et al., 2000)., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Paranthropus aethiopicus The cranium KNM-WT 17000 was examined first-hand for this study., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Homo erectus Samples from Buia, Chemeron, Daka, Dmanisi, East and West Lake Turkana, Gona, Hexian, Konso, Mojokerto, Olduvai Gorge, Sangiran, Swartkrans, Trinil, and Zhoukoudian were included in this study. South African material is of special interest in this comparison because of the geographic proximity, and because of the difficulty of clearly identifying Homo specimens within the large fossil sample from Swartkrans. In particular, the following specimens from Swartkrans are considered to represent H. erectus: SK 15, SK 18a, SK 27, SK 43, SK 45, SK 68, SK 847, SK 878, SK 2635, SKW 3114, SKX 257/258, SKX 267/ 2671, SKX 268, SKX 269, SKX 334, SKX 339, SKX 610, SKX 1756, SKX 2354, SKX 2355, SKX 2356, and SKX 21204. It has been suggested (Grine et al., 1993, 1996) that SK 847 and Stw 53 might represent the same taxon, and that this taxon is a currently undiagnosed species of Homo in South Africa. However, we agree with Clarke (1977; 2008) that SK 847 can be attributed to H. erectus, and that Stw 53 cannot. Because there is no clear indication that more than one species of Homo is represented in the Swartkrans sample, we consider all this material to belong to H. erectus. We considered ‘ Homo ergaster ’ (and also ‘ Homo aff. erectus ’ from Wood, 1991) to be synonyms of H. erectus for this study; Turkana Basin specimens that are attributed to H. erectus thus include KNM-ER 730, KNM-ER 820, KNM-ER 992, KNM- ER 1808, KNM-ER 3733, KNM-ER 3883, KNM-ER 42700, KNM-WT 15000. Olduvai specimens include OH 9, OH 12 and OH 28. Original fossil materials from Chemeron, Lake Turkana, Swartkrans, Trinil, and Dmanisi were examined first-hand by the authors, while the remainder were based on casts and published reports (Weidenreich, 1943; Wood, 1991; Anton´, 2003; Rightmire et al., 2006; Suwa et al., 2007). A large number of postcranial specimens have been collected from the Turkana Basin and appear consistent with the anatomical range otherwise found in Homo, and inconsistent with known samples of Australopithecus and Paranthropus from elsewhere. These include KNM-ER 1472, KNM-ER 1481, KNM- ER 3228, KNM-ER 737, and others. We may add other fossils from other sites lacking association with craniodental material, such as the partial BOU-VP 12/1 skeleton and even the Gona pelvis. These specimens attributable to Homo but not necessarily to a particular species did inform our understanding of variability within the genus, but for the most part these specimens do not inform our differential diagnosis of H. naledi relative to particular species. For example, the key element of femoral morphology of H. naledi in contrast to other species is the presence of two well-defined mediolaterally running pillars in the femoral neck; the isolated specimens of early Homo do not contradict this apparent autapomorphy. Likewise, no isolated specimens inform us about the humanlike aspects of foot morphology in H. naledi. In these cases, the lack of associations for this evidence actually is less important than the lack of specimens that replicate the distinctive features of the H. naledi morphology., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 27, DOI: 10.7554/eLife.09560

Australopithecus garhi The cranium BOU-VP-12/130 from Bouri was included, with data taken from a published report (Asfaw et al., 1999)., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Australopithecus sediba The partial skeletons MH1 and MH2 from Malapa, South Africa were included in this study, based on examination of the original specimens by the authors., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Middle Pleistocene Homo Specimens from the latest Lower Pleistocene and MP of Europe and Africa that cannot be attributed to H. erectus were included in our comparisons. These include fossils that have been attributed to H. heidelbergensis, H. rhodesiensis, ‘archaic H. sapiens ’, or ‘evolved H. erectus ’ by a variety of other authors. Specimens attributed to MP Homo include materials from Eliye Springs, Arago, Atapuerca Sima de los Huesos, Bodo, Broken Hill, Cave of Hearths, Ceprano, Dali, Elandsfontein, Jinniushan, Kapthurin, Mauer, Narmada, Ndutu, Petralona, Reilingen-Schwetzingen, Solo, Steinheim, Swanscombe. This grouping includes the following specimens: KNM-ES 11693, Arago 2, Arago 13, Arago 21, Atapuerca 1, Atapuerca 2, Atapuerca 4, Atapuerca 5, Atapuerca 6, Cave of Hearths, SAM-PQ- EH1, Kabwe, Mauer, Ndutu, Sale´, Petralona, Reilingen-Schwetzingen, Steinheim.

© Berger et al. Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.

Long-term trends in robusticity of lower limb bones in the genus Homo through the Pleistocene until the present have been proposed, which have been interpreted as a consequence of decreasing levels of mobility and activity patterns, changes in lifestyle, and environmental factors. There has also long been evidence that skeletal strength increases over an individual's lifespan. This increase is caused by continuous bone remodeling that optimizes the structure of a bone to resist mechanical loadings and creates a balance between endosteal resorption and subperiosteal apposition. However, none of the previous studies of temporal trends in robusticity has considered both processes and analyzed how individual age-related robusticity might influence higher-level temporal trends. This paper therefore explores temporal trends in robusticity of lower limb long bones within the genus Homo and considers how individual ages-at-death can confound published evolutionary trends, given the fact that some aspects of relative bone strength tend to increase over individual lifespans. Cross-sectional diaphyseal properties of the midshaft and proximal femur and midshaft tibia of Pleistocene and early Holocene individuals, together with data on age-at-death are used to analyze changes in relative bone strength relative to individuals' ages and evolutionary time. The results show increasing bone strength in adulthood until the fourth decade and then a slight decrease, an observation that conforms to previously published results on recent human populations. However, no significant impact of age-at-death on the trends along an evolutionary trajectory has been detected. The evolutionary trends in femoral and tibial relative strength can be described as fluctuating, probably as a consequence of changing mobility patterns, environmentally and technologically influenced behaviors, and demographic processes. The differences between evolutionary trends published in several studies are explained primarily as a result of different ways of standardizing cross-sectional parameters for size, and differences in sample composition.

Dans un article publié dans le n° 20 de la revue Paleo (Madelaine et al. 2008) nous avons annoncé la découverte d’une série de 11 nouveaux restes humains que nous avons pu rapporter à l’individu n° 1 (Vandermeersch et Trinkaus 1995) de Regourdou mis au jour en septembre 1957. Certains, parmi les plus conséquents (fémur, tibia, fibula), représentaient ou complétaient des régions squelettiques supposées manquantes en raison de gestes funéraires (Bonifay et al. 2007).Dans cette contribution, nous augmentons la représentation squelettique de Regourdou 1 avec de nouvelles pièces. Elles proviennent d’une part de la collection de faune (propriété du Musée national de Préhistoire depuis 2002), de celle de la famille Constant et d’autre part, des collections du Musée d’Art et d’Archéologie de Périgueux. Deux ossements, une diaphyse fémorale gauche et un tibia, proviennent des collections de faune du Musée de site de Regourdou. Mais la morphologie du second ne permet pas de le rapporter à l’Homme de Néandertal.Ces nouvelles découvertes nous permettent donc d’enrichir nos connaissances sur la variabilité anatomique néandertalienne et sur l’histoire de l’occupation du site. Elles valident la présence d’au moins un second adulte moustérien, uniquement représenté par un calcaneus droit. Elles nous autorisent aussi à revenir sur une hypothèse intéressant l’histoire taphonomique du sujet le plus complet (Regourdou 1), hypothèse que nous avions avancée en 2008 et qui apparaît désormais fausse. Enfin, l’origine de l’individu représenté par un tibia de morphologie moderne est inconnue. Il nécessitera une tentative de datation absolue directe. In an article published in Paleo number 20 (Madelaine et al. 2008), we announced the discovery of a new series of 11 human remains that we associated with Regourdou individual number 1 (Vandermeersch and Trinkaus 1995), a specimen first brought to light in September 1957. Some of these new bones, in particular the larger ones (femur, tibia, fibula) represent or make more complete skeletal elements that were presumed to have been missing as the result of funerary acts (Bonifay et al. 2007).In this contribution, we increase the skeletal representation of Regourdou 1 with additional new pieces.These come in part from the site’s faunal collections (property of the Musée national de Préhistoire since 2002), but also from the collections of the Musée d’Art et d’Archéologie de Périgueux. In addition, two pieces: a left femoral diaphysis and the proximal half of a tibia come from the collection of the Constant family, but the morphology of the latter does not appear to be that of a Neandertal.These new discoveries increase our knowledge of Neandertal anatomical variability and the history of the site’s occupation, and also permit us to confirm the presence of at least one second adult Mousterian-associated individual, who is thus far represented solely by a right calcaneus. They also allow us to revisit an interesting hypothesis regarding the taphonomic history of the most complete individual from the site (Regourdou 1), a hypothesis we put forth in 2008 that now turns out to be false. Finally, the origin of the individual represented by the tibia (which evinces modern morphology) is unknown. We will therefore need to obtain an absolute direct date on it.