Initial Upper Palaeolithic Homo sapiens from Bacho Kiro Cave, Bulgaria.

The Middle to Upper Palaeolithic transition in Europe witnessed the replacement and partial absorption of local Neanderthal populations by Homo sapiens populations of African origin1. However, this process probably varied across regions and its details remain largely unknown. In particular, the duration of chronological overlap between the two groups is much debated, as are the implications of this overlap for the nature of the biological and cultural interactions between Neanderthals and H. sapiens. Here we report the discovery and direct dating of human remains found in association with Initial Upper Palaeolithic artefacts2, from excavations at Bacho Kiro Cave (Bulgaria). Morphological analysis of a tooth and mitochondrial DNA from several hominin bone fragments, identified through proteomic screening, assign these finds to H. sapiens and link the expansion of Initial Upper Palaeolithic technologies with the spread of H. sapiens into the mid-latitudes of Eurasia before 45 thousand years ago3. The excavations yielded a wealth of bone artefacts, including pendants manufactured from cave bear teeth that are reminiscent of those later produced by the last Neanderthals of western Europe4-6. These finds are consistent with models based on the arrival of multiple waves of H. sapiens into Europe coming into contact with declining Neanderthal populations7,8.

A late Middle Pleistocene Denisovan mandible from the Tibetan Plateau.

Denisovans are members of a hominin group who are currently only known directly from fragmentary fossils, the genomes of which have been studied from a single site, Denisova Cave1-3 in Siberia. They are also known indirectly from their genetic legacy through gene flow into several low-altitude East Asian populations4,5 and high-altitude modern Tibetans6. The lack of morphologically informative Denisovan fossils hinders our ability to connect geographically and temporally dispersed fossil hominins from Asia and to understand in a coherent manner their relation to recent Asian populations. This includes understanding the genetic adaptation of humans to the high-altitude Tibetan Plateau7,8, which was inherited from the Denisovans. Here we report a Denisovan mandible, identified by ancient protein analysis9,10, found on the Tibetan Plateau in Baishiya Karst Cave, Xiahe, Gansu, China. We determine the mandible to be at least 160 thousand years old through U-series dating of an adhering carbonate matrix. The Xiahe specimen provides direct evidence of the Denisovans outside the Altai Mountains and its analysis unique insights into Denisovan mandibular and dental morphology. Our results indicate that archaic hominins occupied the Tibetan Plateau in the Middle Pleistocene epoch and successfully adapted to high-altitude hypoxic environments long before the regional arrival of modern Homo sapiens.

Dental data challenge the ubiquitous presence of Homo in the Cradle of Humankind.

The origins of Homo, as well as the diversity and biogeographic distribution of early Homo species, remain critical outstanding issues in paleoanthropology. Debates about the recognition of early Homo, first appearance dates, and taxonomic diversity within Homo are particularly important for determining the role that southern African taxa may have played in the origins of the genus. The correct identification of Homo remains also has implications for reconstructing phylogenetic relationships between species of Australopithecus and Paranthropus, and the links between early Homo species and Homo erectus. We use microcomputed tomography and landmark-free deformation-based three-dimensional geometric morphometrics to extract taxonomically informative data from the internal structure of postcanine teeth attributed to Early Pleistocene Homo in the southern African hominin-bearing sites of Sterkfontein, Swartkrans, Drimolen, and Kromdraai B. Our results indicate that, from our sample of 23 specimens, only 4 are unambiguously attributed to Homo, 3 of them coming from Swartkrans member 1 (SK 27, SK 847, and SKX 21204) and 1 from Sterkfontein (Sts 9). Three other specimens from Sterkfontein (StW 80 and 81, SE 1508, and StW 669) approximate the Homo condition in terms of overall enamel-dentine junction shape, but retain Australopithecus-like dental traits, and their generic status remains unclear. The other specimens, including SK 15, present a dominant australopith dental signature. In light of these results, previous dietary and ecological interpretations can be reevaluated, showing that the geochemical signal of one tooth from Kromdraai (KB 5223) and two from Swartkrans (SK 96 and SKX 268) is consistent with that of australopiths.

Trabecular architecture of the distal femur in extant hominids.

Extant great apes are characterized by a wide range of locomotor, postural and manipulative behaviours that each require the limbs to be used in different ways. In addition to external bone morphology, comparative investigation of trabecular bone, which (re-)models to reflect loads incurred during life, can provide novel insights into bone functional adaptation. Here, we use canonical holistic morphometric analysis (cHMA) to analyse the trabecular morphology in the distal femoral epiphysis of Homo sapiens (n = 26), Gorilla gorilla (n = 14), Pan troglodytes (n = 15) and Pongo sp. (n = 9). We test two predictions: (1) that differing locomotor behaviours will be reflected in differing trabecular architecture of the distal femur across Homo, Pan, Gorilla and Pongo; (2) that trabecular architecture will significantly differ between male and female Gorilla due to their different levels of arboreality but not between male and female Pan or Homo based on previous studies of locomotor behaviours. Results indicate that trabecular architecture differs among extant great apes based on their locomotor repertoires. The relative bone volume and degree of anisotropy patterns found reflect habitual use of extended knee postures during bipedalism in Homo, and habitual use of flexed knee posture during terrestrial and arboreal locomotion in Pan and Gorilla. Trabecular architecture in Pongo is consistent with a highly mobile knee joint that may vary in posture from extension to full flexion. Within Gorilla, trabecular architecture suggests a different loading of knee in extension/flexion between females and males, but no sex differences were found in Pan or Homo, supporting our predictions. Inter- and intra-specific variation in trabecular architecture of distal femur provides a comparative context to interpret knee postures and, in turn, locomotor behaviours in fossil hominins.

Sex-biased sampling may influence Homo naledi tooth size variation.

A frequent source of debate in paleoanthropology concerns the taxonomic unity of fossil assemblages, with many hominin samples exhibiting elevated levels of variation that can be interpreted as indicating the presence of multiple species. By contrast, the large assemblage of hominin fossils from the Rising Star cave system, assigned to Homo naledi, exhibits a remarkably low degree of variation for most skeletal elements. Many factors can contribute to low sample variation, including genetic drift, strong natural selection, biased sex ratios, and sampling of closely related individuals. In this study, we tested for potential sex-biased sampling in the Rising Star dental sample. We compared coefficients of variation for the H. naledi teeth to those for eight extant hominoid samples. We used a resampling procedure that generated samples from the extant taxa that matched the sample size of the fossil sample for each possible Rising Star dental sex ratio. We found that variation at four H. naledi tooth positions-I2, M1, P4, M1-is so low that the possibility that one sex is represented by few or no individuals in the sample cannot be excluded. Additional evidence is needed to corroborate this inference, such as ancient DNA or enamel proteome data, and our study design does not address other potential factors that would account for low sample variation. Nevertheless, our results highlight the importance of considering the taphonomic history of a hominin assemblage and suggest that sex-biased sampling is a plausible explanation for the low level of phenotypic variation found in some aspects of the current H. naledi assemblage.

Author Correction: New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens.

In the originally published version of this Letter, the x axis in Fig. 3a should have been: 'PC1: 26%' rather than 'PC1: 46%', and the y axis should have been: 'PC2: 16%' rather than 'PC2: 29%'. We also noticed an error in the numbering of the fossils from Qafzeh: Qafzeh 27 should be removed, and Qafzeh 26 is actually Qafzeh 25, following Tillier (2014)1 and Schuh et al. (2017)2 and personal communication with B. Vandermeersch and M. D. Garralda. The correct enumeration of Qafzeh samples in the 'Mandibular metric data' section of the Methods is therefore: 'Qafzeh (9, 25)' rather than 'Qafzeh (9, 26, 27)'. Owing to the removal of Qafzeh 27, the convex hull of early modern humans changes slightly in Extended Data Fig. 1c. The sample sizes in Extended Data Fig. 1c should have read: Middle Pleistocene archaic Homo n = 19 (instead of 11), Neanderthals n = 40 (instead of 41), early modern humans n = 12 (instead of 7), and recent modern humans n = 46 (instead of 48). In Extended Data Table 2, the mean and standard deviation of corpus height and breadth at mental foramen for early modern humans should have been: x̅ = 33.15, σ = 3.26 for height (rather than x̅ = 34.23, σ = 4.57); and x̅ = 16.25, σ = 1.28 for breadth (rather than x̅ = 16.04, σ = 1.75). Accordingly, n = 12 (rather than n = 13) for both breadth and height. These errors have been corrected in the Letter online (the original Extended Data Fig. 1 is shown in Supplementary Information to this Amendment). These changes do not alter any inferences drawn from the data.

Cortical bone distribution of the proximal phalanges in great apes: implications for reconstructing manual behaviours.

Primate fingers are typically in direct contact with the environment during both locomotion and manipulation, and aspects of external phalangeal morphology are known to reflect differences in hand use. Since bone is a living tissue that can adapt in response to loading through life, the internal bone architecture of the manual phalanges should also reflect differences in manual behaviours. Here, we use the R package Morphomap to analyse high-resolution microCT scans of hominid proximal phalanges of digits 2-5 to determine whether cortical bone structure reflects variation in manual behaviours between bipedal (Homo), knuckle-walking (Gorilla, Pan) and suspensory (Pongo) taxa. We test the hypothesis that relative cortical bone distribution patterns and cross-sectional geometric properties will differ both among extant great apes and across the four digits due to locomotor and postural differences. Results indicate that cortical bone structure reflects the varied hand postures employed by each taxon. The phalangeal cortices of Pongo are significantly thinner and have weaker cross-sectional properties relative to the African apes, yet thick cortical bone under their flexor sheath ridges corresponds with predicted loading during flexed finger grips. Knuckle-walking African apes have even thicker cortical bone under the flexor sheath ridges, as well as in the region proximal to the trochlea, but Pan also has thicker diaphyseal cortices than Gorilla. Humans display a distinct pattern of distodorsal thickening, as well as relatively thin cortices, which may reflect the lack of phalangeal curvature combined with frequent use of flexed fingered hand grips during manipulation. Within each taxon, digits 2-5 have a similar cortical distribution in Pongo, Gorilla and, unexpectedly, Homo, which suggest similar loading of all fingers during habitual locomotion or hand use. In Pan, however, cortical thickness differs between the fingers, potentially reflecting differential loading during knuckle-walking. Inter- and intra-generic variation in phalangeal cortical bone structure reflects differences in manual behaviours, offering a comparative framework for reconstructing hand use in fossil hominins.

Descriptive catalog of Homo naledi dental remains from the 2013 to 2015 excavations of the Dinaledi Chamber, site U.W. 101, within the Rising Star cave system, South Africa.

More than 150 hominin teeth, dated to ∼330-241 thousand years ago, were recovered during the 2013-2015 excavations of the Dinaledi Chamber of the Rising Star cave system, South Africa. These fossils comprise the first large single-site sample of hominin teeth from the Middle Pleistocene of Africa. Though scattered remains attributable to Homo sapiens, or their possible lineal ancestors, are known from older and younger sites across the continent, the distinctive morphological feature set of the Dinaledi teeth supports the recognition of a novel hominin species, Homo naledi. This material provides evidence of African Homo lineage diversity that lasts until at least the Middle Pleistocene. Here, a catalog, anatomical descriptions, and details of preservation and taphonomic alteration are provided for the Dinaledi teeth. Where possible, provisional associations among teeth are also proposed. To facilitate future research, we also provide access to a catalog of surface files of the Rising Star jaws and teeth.

A tooth crown morphology framework for interpreting the diversity of primate dentitions.

Variation in tooth crown morphology plays a crucial role in species diagnoses, phylogenetic inference, and the reconstruction of the evolutionary history of the primate clade. While a growing number of studies have identified developmental mechanisms linked to tooth size and cusp patterning in mammalian crown morphology, it is unclear (1) to what degree these are applicable across primates and (2) which additional developmental mechanisms should be recognized as playing important roles in odontogenesis. From detailed observations of lower molar enamel-dentine junction morphology from taxa representing the major primate clades, we outline multiple phylogenetic and developmental components responsible for crown patterning, and formulate a tooth crown morphology framework for the holistic interpretation of primate crown morphology. We suggest that adopting this framework is crucial for the characterization of tooth morphology in studies of dental development, discrete trait analysis, and systematics.

New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens.

Fossil evidence points to an African origin of Homo sapiens from a group called either H. heidelbergensis or H. rhodesiensis. However, the exact place and time of emergence of H. sapiens remain obscure because the fossil record is scarce and the chronological age of many key specimens remains uncertain. In particular, it is unclear whether the present day 'modern' morphology rapidly emerged approximately 200 thousand years ago (ka) among earlier representatives of H. sapiens or evolved gradually over the last 400 thousand years. Here we report newly discovered human fossils from Jebel Irhoud, Morocco, and interpret the affinities of the hominins from this site with other archaic and recent human groups. We identified a mosaic of features including facial, mandibular and dental morphology that aligns the Jebel Irhoud material with early or recent anatomically modern humans and more primitive neurocranial and endocranial morphology. In combination with an age of 315 ± 34 thousand years (as determined by thermoluminescence dating), this evidence makes Jebel Irhoud the oldest and richest African Middle Stone Age hominin site that documents early stages of the H. sapiens clade in which key features of modern morphology were established. Furthermore, it shows that the evolutionary processes behind the emergence of H. sapiens involved the whole African continent.

Evidence for habitual climbing in a Pleistocene hominin in South Africa.

Bipedalism is a defining trait of the hominin lineage, associated with a transition from a more arboreal to a more terrestrial environment. While there is debate about when modern human-like bipedalism first appeared in hominins, all known South African hominins show morphological adaptations to bipedalism, suggesting that this was their predominant mode of locomotion. Here we present evidence that hominins preserved in the Sterkfontein Caves practiced two different locomotor repertoires. The trabecular structure of a proximal femur (StW 522) attributed to Australopithecus africanus exhibits a modern human-like bipedal locomotor pattern, while that of a geologically younger specimen (StW 311) attributed to either Homo sp. or Paranthropus robustus exhibits a pattern more similar to nonhuman apes, potentially suggesting regular bouts of both climbing and terrestrial bipedalism. Our results demonstrate distinct morphological differences, linked to behavioral differences between Australopithecus and later hominins in South Africa and contribute to the increasing evidence of locomotor diversity within the hominin clade.

Ontogenetic changes to metacarpal trabecular bone structure in mountain and western lowland gorillas.

The trabecular bone morphology of adult extant primates has been shown to reflect mechanical loading related to locomotion. However, ontogenetic studies of humans and other mammals suggest an adaptive lag between trabecular bone response and current mechanical loading patterns that could result in adult trabecular bone morphology reflecting juvenile behaviours. This study investigates ontogenetic changes in the trabecular bone structure of the third metacarpal of mountain gorillas (Gorilla beringei beringei; n = 26) and western lowland gorillas (Gorilla gorilla gorilla; n = 26) and its relationship to expected changes in locomotor loading patterns. Results show that trabecular bone reflects predicted mechanical loading throughout ontogeny. Bone volume fraction, trabecular thickness and trabecular number are low at birth and increase with age, although degree of anisotropy remains relatively stable throughout ontogeny. A high concentration of bone volume fraction can be observed in the distopalmar region of the third metacarpal epiphysis in early ontogeny, consistent with the high frequency of climbing, suspensory and other grasping behaviours in young gorillas. High trabecular bone concentration increases dorsally in the epiphysis during the juvenile period as terrestrial knuckle-walking becomes the primary form of locomotion. However, fusion of the epiphysis does not take place until 10-11 years of age, and overall trabecular structure does not fully reflect the adult pattern until 12 years of age, indicating a lag between adult-like behaviours and adult-like trabecular morphology. We found minimal differences in trabecular ontogeny between mountain and western lowland gorillas, despite presumed variation in the frequencies of arboreal locomotor behaviours. Altogether, ontogenetic changes in Gorilla metacarpal trabecular structure reflect overall genus-level changes in locomotor behaviours throughout development, but with some ontogenetic lag that should be considered when drawing functional conclusions from bone structure in extant or fossil adolescent specimens.

Calcar femorale variation in extant and fossil hominids: Implications for identifying bipedal locomotion in fossil hominins.

The calcar femorale is an internal bony structure of the proximal femur considered to be functionally related to bipedal locomotion. Among extant primates, the presence of a calcar femorale has been so far documented in extant humans and Pan and, among extinct hominins, in the Late Miocene Orrorin, in a Pliocene Australopithecus, and in a Middle Pleistocene Homo specimen. Using high-resolution microcomputed tomography, we investigated the occurrence and morphology (i.e., shape, location, and size) of the calcar femorale in an adult sample of extant humans, Pan troglodytes, Gorilla gorilla, Pongo sp., and Papio ursinus. We also investigated for the first time the occurrence and morphology of a calcar femorale in the adult proximal femoral remains of a Late Miocene great ape (Rudapithecus) and five Plio-Pleistocene hominins from Southern and Eastern Africa (Australopithecus and Paranthropus). We took four measurements: periosteal-to-tip maximum length, maximum length excluding cortical thickness, maximum vertical height, and the distance between the most anterior and posterior limits of the root. To allow for intergeneric comparisons, estimated body size was used to standardize all measurements. Nine of 10 extant humans have a well-developed calcar femorale. Among the African apes, 6 of 10 Pan and 6 of 10 Gorilla also show a distinct calcar femorale. In Pongo (n = 9), it is only present in one captive individual. None of the five investigated Papio specimens show any trace of this structure. Only calcar femorale height, which is systematically taller and extends into the lower part of the lesser trochanter, discriminates humans from extant great apes, except for one Gorilla. The calcar femorale was absent in one Paranthropus robustus and variably developed in all other investigated fossils. These results indicate that this structure cannot be considered as a diagnostic feature of habitual bipedal locomotion and emphasize the need for further investigations of its functional role.

A simple rule governs the evolution and development of hominin tooth size.

The variation in molar tooth size in humans and our closest relatives (hominins) has strongly influenced our view of human evolution. The reduction in overall size and disproportionate decrease in third molar size have been noted for over a century, and have been attributed to reduced selection for large dentitions owing to changes in diet or the acquisition of cooking. The systematic pattern of size variation along the tooth row has been described as a 'morphogenetic gradient' in mammal, and more specifically hominin, teeth since Butler and Dahlberg. However, the underlying controls of tooth size have not been well understood, with hypotheses ranging from morphogenetic fields to the clone theory. In this study we address the following question: are there rules that govern how hominin tooth size evolves? Here we propose that the inhibitory cascade, an activator-inhibitor mechanism that affects relative tooth size in mammals, produces the default pattern of tooth sizes for all lower primary postcanine teeth (deciduous premolars and permanent molars) in hominins. This configuration is also equivalent to a morphogenetic gradient, finally pointing to a mechanism that can generate this gradient. The pattern of tooth size remains constant with absolute size in australopiths (including Ardipithecus, Australopithecus and Paranthropus). However, in species of Homo, including modern humans, there is a tight link between tooth proportions and absolute size such that a single developmental parameter can explain both the relative and absolute sizes of primary postcanine teeth. On the basis of the relationship of inhibitory cascade patterning with size, we can use the size at one tooth position to predict the sizes of the remaining four primary postcanine teeth in the row for hominins. Our study provides a development-based expectation to examine the evolution of the unique proportions of human teeth.

Cortical and trabecular bone structure of the hominoid capitate.

Morphological variation in the hominoid capitate has been linked to differences in habitual locomotor activity due to its importance in movement and load transfer at the midcarpal joint proximally and carpometacarpal joints distally. Although the shape of bones and their articulations are linked to joint mobility, the internal structure of bones has been shown experimentally to reflect, at least in part, the loading direction and magnitude experienced by the bone. To date, it is uncertain whether locomotor differences among hominoids are reflected in the bone microarchitecture of the capitate. Here, we apply a whole-bone methodology to quantify the cortical and trabecular architecture (separately and combined) of the capitate across bipedal (modern Homo sapiens), knuckle-walking (Pan paniscus, Pan troglodytes, Gorilla sp.), and suspensory (Pongo sp.) hominoids (n = 69). It is hypothesized that variation in bone microarchitecture will differentiate these locomotor groups, reflecting differences in habitual postures and presumed loading force and direction. Additionally, it is hypothesized that trabecular and cortical architecture in the proximal and distal regions, as a result of being part of mechanically divergent joints proximally and distally, will differ across these portions of the capitate. Results indicate that the capitate of knuckle-walking and suspensory hominoids is differentiated from bipedal Homo primarily by significantly thicker distal cortical bone. Knuckle-walking taxa are further differentiated from suspensory and bipedal taxa by more isotropic trabeculae in the proximal capitate. An allometric analysis indicates that size is not a significant determinate of bone variation across hominoids, although sexual dimorphism may influence some parameters within Gorilla. Results suggest that internal trabecular and cortical bone is subjected to different forces and functional adaptation responses across the capitate (and possibly other short bones). Additionally, while separating trabecular and cortical bone is normal protocol of current whole-bone methodologies, this study shows that when applied to carpals, removing or studying the cortical bone separately potentially obfuscates functionally relevant signals in bone structure.

The morphology of the Late Pleistocene hominin remains from the site of La Cotte de St Brelade, Jersey (Channel Islands).

Thirteen permanent fully erupted teeth were excavated at the Paleolithic site of La Cotte de St Brelade in Jersey in 1910 and 1911. These were all found in the same location, on a ledge behind a hearth in a Mousterian occupation level. They were originally identified as being Neanderthal. A fragment of occipital bone was found in a separate locality in a later season. Recent dating of adjacent sediments gives a probable age of <48 ka. The purpose of this article is to provide an updated description of the morphology of this material and consider its likely taxonomic assignment from comparison with Neanderthal and Homo sapiens samples. One of the original teeth has been lost, and we identify one as nonhominin. At least two adult individuals are represented. Cervix shape and the absence of common Neanderthal traits in several teeth suggest affinities with H. sapiens in both individuals, while crown and root dimensions and root morphology of all the teeth are entirely consistent with a Neanderthal attribution, pointing toward a possible shared Neanderthal and H. sapiens ancestry (the likely date of this material corresponds with the time in which both Neanderthals and H. sapiens were present in Europe). The occipital fragment is stratigraphically more recent and does not exhibit any diagnostic Neanderthal features.

Maxillary molar enamel thickness of Plio-Pleistocene hominins.

Enamel thickness remains an important morphological character in hominin systematics and is regularly incorporated into dietary reconstructions in hominin species. We expand upon a previous study of enamel thickness in mandibular molars by examining a large maxillary molar sample of Plio-Pleistocene hominins (n = 62) and a comparative sample of extant nonhuman apes (n = 48) and modern humans (n = 29). 2D mesial planes of section were generated through microtomography, and standard dental tissue variables were measured to calculate average enamel thickness (AET) and relative enamel thickness (RET). AET was also examined across the lingual, occlusal, and buccal regions of the crown. This study confirms previous findings of increasing enamel thickness throughout the Plio-Pleistocene, being thinnest in Australopithecus anamensis and peaking in Australopithecus boisei, with early Homo specimens, exhibiting intermediate enamel thickness. Agreeing with previous findings, 2D plane of section enamel thickness is found to be a poor taxonomic discriminator, with no statistically significant differences observed between fossil hominins. For fossil hominins, modern humans, and Pongo, the occlusal region of enamel was the thickest, and the lingual enamel thickness was greater than buccal. Pan and Gorilla present the opposite pattern with enamel being thinnest occlusally. Comparison at each molar position between the maxilla and mandible revealed very few significant differences in fossil hominins but some evidence of significantly thicker maxillary enamel (AET) in modern humans and thinner maxillary enamel in Pan (RET).

Hominin dental remains from the Pliocene localities at Lomekwi, Kenya (1982-2009).

Increasing evidence for both taxonomic diversity and early stone manufacture during the Pliocene highlights the importance of the hominin fossil record from this epoch in eastern Africa. Here, we describe dental remains from Lomekwi (West Turkana, Kenya), which date from between 3.2 and 3.5 Ma. The sample was collected between 1982 and 2009 and includes five gnathic specimens and a total of 67 teeth (mostly isolated permanent postcanine teeth). Standard linear dimensions indicate that, although the Lomekwi teeth are relatively small, there is broad overlap in size with contemporary Australopithecus afarensis and Australopithecus deyiremeda specimens at most tooth positions. However, some dental characters differentiate this sample from these species, including a relatively large P4 and M3 compared with the M1, a high incidence of well-developed protostylids, and specific accessory molar cuspules. Owing to a lack of well-preserved tooth crowns (and the complete absence of mandibular teeth) in the holotype and paratype of Kenyanthropus platyops, and limited comparable gnathic morphology in the new specimens, it cannot be determined whether these Lomekwi specimens should be attributed to this species. Attribution of these specimens is further complicated by a lack of certainty about position along the tooth row of many of the molar specimens. More comprehensive shape analyses of the external and internal morphology of these specimens, and additional fossil finds, would facilitate the taxonomic attribution of specimens in this taxonomically diverse period of human evolution.

The Neanderthal teeth from Marillac (Charente, Southwestern France): Morphology, comparisons and paleobiology.

Few European sites have yielded human dental remains safely dated to the end of MIS 4/beginning of MIS 3. One of those sites is Marillac (Southwestern France), a collapsed karstic cave where archeological excavations (1967-1980) conducted by B. Vandermeersch unearthed numerous faunal and human remains, as well as a few Mousterian Quina tools. The Marillac sinkhole was occasionally used by humans to process the carcasses of different prey, but there is no evidence for a residential use of the site, nor have any hearths been found. Rare carnivore bones were also discovered, demonstrating that the sinkhole was seasonally used, not only by Neanderthals, but also by predators across several millennia. The lithostratigraphic units containing the human remains were dated to ∼60 kyr. The fossils consisted of numerous fragments of skulls and jaws, isolated teeth and several post-cranial bones, many of them with traces of perimortem manipulations. For those already published, their morphological characteristics and chronostratigraphic context allowed their attribution to Neanderthals. This paper analyzes sixteen unpublished human teeth (fourteen permanent and two deciduous) by investigating the external morphology and metrical variation with respect to other Neanderthal remains and a sample from modern populations. We also investigate their enamel thickness distribution in 2D and 3D, the enamel-dentine junction morphology (using geometric morphometrics) of one molar and two premolars, the roots and the possible expression of taurodontism, as well as pathologies and developmental defects. The anterior tooth use and paramasticatory activities are also discussed. Morphological and structural alterations were found on several teeth, and interpreted in light of human behavior (tooth-pick) and carnivores' actions (partial digestion). The data are interpreted in the context of the available information for the Eurasian Neanderthals.

Trabecular variation in the first metacarpal and manipulation in hominids.

OBJECTIVES: The dexterity of fossil hominins is often inferred by assessing the comparative manual anatomy and behaviors of extant hominids, with a focus on the thumb. The aim of this study is to test whether trabecular structure is consistent with what is currently known about habitually loaded thumb postures across extant hominids. MATERIALS AND METHODS: We analyze first metacarpal (Mc1) subarticular trabecular architecture in humans (Homo sapiens, n = 10), bonobos (Pan paniscus, n = 10), chimpanzees (Pan troglodytes, n = 11), as well as for the first time, gorillas (Gorilla gorilla gorilla, n = 10) and orangutans (Pongo sp., n = 1, Pongo abelii, n = 3 and Pongo pygmaeus, n = 5). Using a combination of subarticular and whole-epiphysis approaches, we test for significant differences in relative trabecular bone volume (RBV/TV) and degree of anisotropy (DA) between species. RESULTS: Humans have significantly greater RBV/TV on the radiopalmar aspects of both the proximal and distal Mc1 subarticular surfaces and greater DA throughout the Mc1 head than other hominids. Nonhuman great apes have greatest RBV/TV on the ulnar aspect of the Mc1 head and the palmar aspect of the Mc1 base. Gorillas possessed significantly lower DA in the Mc1 head than any other taxon in our sample. DISCUSSION: These results are consistent with abduction of the thumb during forceful "pad-to-pad" precision grips in humans and, in nonhuman great apes, a habitually adducted thumb that is typically used in precision and power grips. This comparative context will help infer habitual manipulative and locomotor grips in fossil hominins.