Jean-Jacques Hublin

Most of the Middle Palaeolithic evidence of Central Italy still lacks a reliable chrono-cultural framework mainly due to research history. In this context Grotta dei Santi, a wide cave located on Monte Argentario, on the southern coast of Tuscany, is particularly relevant as it contains a very well preserved sequence including several Mousterian layers. Research carried out at this site in the last years (2007e2017) allowed for a preliminary estimation of its chronology based on a set of radiometric determinations which place the investigated sequence in the time interval between 50 and 40 ka BP. Alongside the chronological issue, this paper mainly focuses on the geoarchaeological and zooarchaeological (micro and macro fauna) studies carried out on the materials retrieved during the 2007e2014 excavation fieldworks. The results of these studies are consistent with those from the radiometric chronology. A state of art concerning the MIS3 Italian sites is also provided in order to highlight the key role Grotta dei Santi may play in the assessment of late Neandertals' behaviour within the framework of the Middle to Upper Palaeolithic transition of Central Italy.

Although it has previously been shown that Neanderthals contributed DNA to modern humans 1,2 , not much is known about the genetic diversity of Neanderthals or the relationship between late Neanderthal populations at the time at which their last interactions with early modern humans occurred and before they eventually disappeared. Our ability to retrieve DNA from a larger number of Neanderthal individuals has been limited by poor preservation of endogenous DNA 3 and contamination of Neanderthal skeletal remains by large amounts of microbial and present-day human DNA 3-5. Here we use hypochlorite treatment 6 of as little as 9 mg of bone or tooth powder to generate between 1-and 2.7-fold genomic coverage of five Neanderthals who lived around 39,000 to 47,000 years ago (that is, late Neanderthals), thereby doubling the number of Neanderthals for which genome sequences are available. Genetic similarity among late Neanderthals is well predicted by their geographical location, and comparison to the genome of an older Neanderthal from the Caucasus 2,7 indicates that a population turnover is likely to have occurred, either in the Caucasus or throughout Europe, towards the end of Neanderthal history. We find that the bulk of Neanderthal gene flow into early modern humans originated from one or more source populations that diverged from the Neanderthals that were studied here at least 70,000 years ago, but after they split from a previously sequenced Neanderthal from Siberia 2 around 150,000 years ago. Although four of the Neanderthals studied here postdate the putative arrival of early modern humans into Europe, we do not detect any recent gene flow from early modern humans in their ancestry.

Recent studies have shown that the use of aquatic resources has greater antiquity in hominin diets than previously thought. At present, it is unclear when hominins started to habitually consume marine resources. This study examines shellfish exploitation from a behavioural ecology perspective, addressing how and when past hunter-gatherers from the Levant used coastal resources for subsistence purposes. We investigate the seasonality of shellfish exploitation in the Levantine Upper Palaeolithic through oxygen isotope analysis on shells of the intertidal rocky shore mollusc Phorcus (Osilinus) turbinatus from the key site Ksâr 'Akil (Lebanon). At this rockshelter, multi-layered archaeological deposits contained remains of both marine and terrestrial molluscs in relatively large quantities, which were consumed and used as tools and ornaments by the occupants of the site. Our results indicate that at the start of the Initial Upper Palaeolithic (IUP), there is no evidence for shellfish consumption. Humans started to take fresh shellfish to the rockshelter from the second half of the IUP onward, albeit in low quantities. During the Early Upper Palaeolithic (EUP) shellfish exploitation became increasingly frequent. Oxygen isotope data show that shellfish exploitation was practised in every season throughout most of the Upper Palaeolithic (UP), with an emphasis on the colder months. This suggests that coastal resources had a central role in early UP foraging strategies, rather than a seasonally restricted supplementary one. Year-round shellfish gathering, in turn, suggests that humans occupied the rockshelter at different times of the year, although not necessarily continuously. Our oxygen isotope data is complemented with broader-scale exploitation patterns of faunal resources, both vertebrate and invertebrate, at the site. The inclusion of coastal marine resources signifies a diversification of the human diet from the EUP onward, which is also observed in foraging practices linked to the exploitation of terrestrial fauna.

When first described, the small calvaria KNM-ER 42700 from Ileret, Kenya, was considered a late juvenile or young adult and assigned to Homo erectus. However, this species attribution has subsequently been challenged because the specimen's neurocranial shape differs substantially from that of H. erectus adults. Here, (1) we describe the postmortem damage and deformation that could have influenced previous shape analyses, (2) present digital reconstructions based on computed tomographic scans correcting for these taphonomic defects, and (3) analyze the reconstructed endocranial shape and form, considering both static allometry among adults and ontogenetic allometry. To this end, we use geometric morpho-metrics to analyze the shape of digital endocasts based on landmarks and semilandmarks. Corroborating previous studies of the external surface, we find that the endocranial shape of KNM-ER 42700 falls outside the known adult variation of H. erectus. With an endocranial volume estimate between 721 and 744 ml, size cannot explain its atypical endocranial shape when static allometry within H. erectus is considered. However, the analysis of ontogenetic allometry suggests that it may be a H. erectus individual that is younger than previously thought and had not yet reached adult endocranial shape. Future work should therefore comprehensively review all cranial indicators of its developmental age, including closure of the spheno-occipital synchondrosis. An alternative hypothesis is that KNM-ER 42700 represents an as yet unidentified species of early Homo. Importantly, KNM-ER 42700 should not be included in the adult hypodigm of H. erectus.

The ecology of Neanderthals is a pressing question in the study of hominin evolution. Diet appears to have played a prominent role in their adaptation to Eurasia. Based on isotope and zooarchaeological studies, Neanderthal diet has been reconstructed as heavily meat-based and generally similar across different environments. This image persists, despite recent studies suggesting more plant use and more variation. However, we have only a fragmentary picture of their dietary ecology, and how it may have varied among habitats, because we lack broad and environmentally representative information about their use of plants and other foods. To address the problem, we examined the plant microremains in Neanderthal dental calculus from five archaeological sites representing a variety of environments from the northern Balkans, and the western, central and eastern Mediterranean. The recovered microremains revealed the consumption of a variety of non-animal foods, including starchy plants. Using a modeling approach, we explored the relationships among microremains and environment, while controlling for chronology. In the process, we compared the effectiveness of various diversity metrics and their shortcomings for studying microbotanical remains, which are often morphologically redundant for identification. We developed Minimum Botanical Units as a new way of estimating how many plant types or parts are present in a microbotanical sample. In contrast to some previous work, we found no evidence that plant use is confined to the southern-most areas of Neanderthal distribution. Although interpreting the ecogeographic variation is limited by the incomplete preservation of dietary micro-remains, it is clear that plant exploitation was a widespread and deeply rooted Neanderthal subsistence strategy, even if they were predominately game hunters. Given the limited dietary variation across Neanderthal range in time and space in both plant and animal food exploitation, we argue that vegetal consumption was a feature of a generally static dietary niche.

Research Cite this article: Wroe S et al. 2018 Computer simulations show that Neanderthal facial morphology represents adaptation to cold and high energy demands, but not heavy biting. Proc. R. Soc. B 285: 20180085. http://dx. Three adaptive hypotheses have been forwarded to explain the distinctive Neanderthal face: (i) an improved ability to accommodate high anterior bite forces, (ii) more effective conditioning of cold and/or dry air and, (iii) adaptation to facilitate greater ventilatory demands. We test these hypotheses using three-dimensional models of Neanderthals, modern humans, and a close outgroup (Homo heidelbergensis), applying finite-element analysis (FEA) and computational fluid dynamics (CFD). This is the most comprehensive application of either approach applied to date and the first to include both. FEA reveals few differences between H. heidel-bergensis, modern humans, and Neanderthals in their capacities to sustain high anterior tooth loadings. CFD shows that the nasal cavities of Nean-derthals and especially modern humans condition air more efficiently than does that of H. heidelbergensis, suggesting that both evolved to better withstand cold and/or dry climates than less derived Homo. We further find that Neanderthals could move considerably more air through the nasal pathway than could H. heidelbergensis or modern humans, consistent with the propositions that, relative to our outgroup Homo, Neanderthal facial morphology evolved to reflect improved capacities to better condition cold, dry air, and, to move greater air volumes in response to higher energetic requirements.

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 1 or evolved gradually over the last 400 thousand years 2. 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) 3 , 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.

The timing and location of the emergence of our species and of associated behavioural changes are crucial for our understanding of human evolution. The earliest fossil attributed to a modern form of Homo sapiens comes from eastern Africa and is approximately 195 thousand years old 1,2 , therefore the emergence of modern human biology is commonly placed at around 200 thousand years ago 3,4. The earliest Middle Stone Age assemblages come from eastern and southern Africa but date much earlier 5–7. Here we report the ages, determined by thermoluminescence dating, of fire-heated flint artefacts obtained from new excavations at the Middle Stone Age site of Jebel Irhoud, Morocco, which are directly associated with newly discovered remains of H. sapiens 8. A weighted average age places these Middle Stone Age artefacts and fossils at 315 ± 34 thousand years ago. Support is obtained through the recalculated uranium series with electron spin resonance date of 286 ± 32 thousand years ago for a tooth from the Irhoud 3 hominin mandible. These ages are also consistent with the faunal and microfaunal 9 assemblages and almost double the previous age estimates for the lower part of the deposits 10,11. The north African site of Jebel Irhoud contains one of the earliest directly dated Middle Stone Age assemblages, and its associated human remains are the oldest reported for H. sapiens. The emergence of our species and of the Middle Stone Age appear to be close in time, and these data suggest a larger scale, potentially pan-African, origin for both.

The presence of Neandertals in Europe and Western Eurasia before the arrival of anatomically modern humans is well supported by archaeological and paleontological data. In contrast, fossil evidence for Denisovans, a sister group of Neandertals recently identified on the basis of DNA sequences, is limited to three specimens, all of which originate from Denisova Cave in the Altai Mountains (Siberia, Russia). We report the retrieval of DNA from a deciduous lower second molar (Denisova 2), discovered in a deep stratigraphic layer in Denisova Cave, and show that this tooth comes from a female Denisovan individual. On the basis of the number of " missing substitutions " in the mitochondrial DNA determined from the specimen, we find that Denisova 2 is substantially older than two of the other Denisovans, reinforcing the view that Denisovans were likely to have been present in the vicinity of Denisova Cave over an extended time period. We show that the level of nuclear DNA sequence diversity found among Denisovans is within the lower range of that of present-day human populations.

The late Middle Palaeolithic (MP) settlement patterns in the Levant included the repeated use of caves and open landscape sites. The fossil record shows that two types of hominins occupied the region during this period—Neandertals and Homo sapiens. Until recently, diagnostic fossil remains were found only at cave sites. Because the two populations in this region left similar material cultural remains, it was impossible to attribute any open-air site to either species. In this study, we present newly discovered fossil remains from intact archaeological layers of the open-air site 'Ein Qashish, in northern Israel. The hominin remains represent three individuals: EQH1, a nondiagnostic skull fragment; EQH2, an upper right third molar (RM 3); and EQH3, lower limb bones of a young Neandertal male. EQH2 and EQH3 constitute the first diagnostic anatomical remains of Neandertals at an open-air site in the Levant. The optically stimulated luminescence ages suggest that Neandertals repeatedly visited 'Ein Qashish between 70 and 60 ka. The discovery of Neandertals at open-air sites during the late MP reinforces the view that Neandertals were a resilient population in the Levant shortly before Upper Palaeolithic Homo sapiens populated the region. The Middle Palaeolithic (MP) of the southern Levant is a significant period for the study of human evolution because two types of hominins, Neandertals and Homo sapiens, occupied the region at that time (see, for example, refs 1 and 2). Diagnostic fossil remains of the two species have been found in the Mediterranean woodland region, but until recently, they were discovered only at cave sites (Fig. 1). The absolute chronology of the Levantine MP fossils indicates that H. sapiens existed there between 120 and 90 ka and again from 55 ka on; Neandertals existed in that region between ca. 80 and ca. 55 ka 3–16. The genomic evidence suggests gene flow from early H. sapiens to

Sub-fossilised remains may still contain highly degraded ancient DNA (aDNA) useful for palaeogenetic investigations. Whether X-ray computed [micro-] tomography ([µ]CT) imaging of these fossils may further damage aDNA remains debated. Although the effect of X-ray on DNA in living organisms is well documented, its impact on aDNA molecules is unexplored. Here we investigate the effects of synchrotron X-ray irradiation on aDNA from Pleistocene bones. A clear correlation appears between decreasing aDNA quantities and accumulating X-ray dose-levels above 2000 Gray (Gy). We further find that strong X-ray irradiation reduces the amount of nucleotide misincorporations at the aDNA molecule ends. No representative effect can be detected for doses below 200 Gy. Dosimetry shows that conventional μ CT usually does not reach the risky dose level, while classical synchrotron imaging can degrade aDNA significantly. Optimised synchrotron protocols and simple rules introduced here are sufficient to ensure that fossils can be scanned without impairing future aDNA studies. peer-reviewed)

The diminutive middle ear ossicles (malleus, incus, stapes) housed in the tympanic cavity of the temporal bone play an important role in audition. The few known ossicles of Neandertals are distinctly different from those of anatomically modern humans (AMHs), despite the close relationship between both human species. Although not mutually exclusive, these differences may affect hearing capacity or could reflect covariation with the surrounding temporal bone. Until now, detailed comparisons were hampered by the small sample of Neandertal ossicles and the unavailability of methods combining analyses of ossicles with surrounding structures. Here, we present an analysis of the largest sample of Neandertal ossicles to date, including many previously unknown specimens, covering a wide geographic and temporal range. Microcomputed tomography scans and 3D geometric morphometrics were used to quantify shape and functional properties of the ossicles and the tympanic cavity and make comparisons with recent and extinct AMHs as well as African apes. We find striking morphological differences between ossicles of AMHs and Neandertals. Ossicles of both Neandertals and AMHs appear derived compared with the inferred ancestral morphology, albeit in different ways. Brain size increase evolved separately in AMHs and Neandertals, leading to differences in the tympanic cavity and, consequently, the shape and spatial configuration of the ossicles. Despite these different evolutionary trajectories, functional properties of the middle ear of AMHs and Neandertals are largely similar. The relevance of these functionally equivalent solutions is likely to conserve a similar auditory sensitivity level inherited from their last common ancestor. middle ear | homo | 3D shape | covariation T he hominin fossil record can only provide indirect information about auditory capacities of our extinct relatives. Inferences about the evolution of the human sense of hearing require understanding of the interplay between form and function in extant species. When auditory capacities are visualized as audiograms, plotting the sensitivity for different frequencies, anatomically modern humans (AMHs) differ from the W-shaped pattern found in most anthropoid primates. AMHs are characterized by a drastically lowered high-frequency cutoff and maintaining high sensitivity in the low to midfrequencies, resulting in a U-shaped audiogram (1–7). In primates, such hearing variability is assumed to be partly related to forms of vocalization and habitat acoustics (8–10). Diverse hearing capabilities are also related to the morphology of the diminutive middle ear ossicles housed in the tympanic cavity (11, 12). The malleus, incus, and stapes form the ossicular chain that connects the tympanic membrane to the oval window of the inner ear. These bones play an important role in audition by amplifying and regulating the sound waves transmitted to the cochlea (11, 13–15). In particular, the middle ear acts as a transformer that matches the impedances between the air and the perilymph of the cochlea (16), participating in the tuning of the sensitivity levels. Recent analyses have emphasized a distinctly derived morphology of the ossicles of AMHs compared with extant great apes (17, 18), suggesting that the ossicles of extinct hominins may provide insights into the origin of the distinct auditory capacities of AMHs. Until recently, only a few isolated Neandertal ossicles were known, and their morphology differs consistently from AMH ossicles (19– 25). Because the external acoustic meatus and cochlea have nearly identical dimensions in AMHs and Neandertals (26–28), such shape differences of the ossicles could indicate differences in auditory capacities and with it, potential implications for habitat preference and aspects of vocal communication. However, the temporal bone housing the ossicles is well-known to differentiate Neandertals from AMHs, and some of its structures express morphological covariation (29, 30). Therefore, differences in ossicle morphology between Neandertals and AMHs could also reflect variation in the spatial relationship of the ossicles within the tympanic cavity because of differences in the placement of either the oval window or the tympanic membrane (24). Comparative and functional investigations of Neandertal os-sicles were previously limited by the small sample size. High-resolution computed tomography (CT) has made it possible to study the morphology of ossicles trapped in the tympanic cavity. Here, we analyze the largest sample of Neandertal ossicles to date (22 ossicles from 14 individuals), covering a wide geographic and temporal range. We compare these fossils with Holocene and Pleistocene AMHs and African apes and investigate how previously observed ossicle characteristics fit into the Neandertal bauplan and how it evolved. Studying ossicles is methodologically challenging because of their small size and complex 3D shapes. To quantify ossicle shape, we apply a 3D geometric morphometric approach (18). We also test how differences in ossicle morphology affect the impedance matching function of the middle ear. As a functional measure, we chose an ideal transformer ratio (ITR), namely the pressure gain, which is the product of the area ratio between the tympanic membrane and stapes footplate and the lever ratio of the malleus and incus functional lengths (31–33). Finally, we analyze how ossicle shape Significance Middle ear ossicles are critical for audition and rarely preserved in fossils. Based on microcomputed tomography images, our comparative 3D shape analysis of Neandertal ossicles shows striking shape differences between Neandertals and anatomically modern humans (AMHs). However, these morphological differences do not affect the functional properties of the ossicles, potentially indicating consistent aspects of vocal communication in Nean-dertals and AMHs. Instead, a strong relationship between ossicle morphology and tympanic cavity architecture is found.

Archaeological sites in northern Africa provide a rich record of increasing importance for the origins of modern human behaviour and for understanding human dispersal out of Africa. However, the timing and nature of Palaeolithic human behaviour and dispersal across northwestern Africa (the Maghreb), and their relationship to local environmental conditions, remain poorly understood. The cave of Rhafas (northeast Morocco) provides valuable chronological information about cultural changes in the Maghreb during the Palaeolithic due to its long stratified archaeological sequence comprising Middle Stone Age (MSA), Later Stone Age (LSA) and Neolithic occupation layers. In this study, we apply optically stimulated luminescence (OSL) dating on sand-sized quartz grains to the cave deposits of Rha-fas, as well as to a recently excavated section on the terrace in front of the cave entrance. We hereby provide a revised chronostratigraphy for the archaeological sequence at the site. We combine these results with geological and sedimentological multi-proxy investigations to gain insights into site formation processes and the palaeoenvironmental record of the region. The older sedimentological units at Rhafas were deposited between 135 ka and 57 ka (MIS 6 –MIS 3) and are associated with the MSA technocomplex. Tanged pieces start to occur in the archaeological layers around 109 ka, which is consistent with previously published chronological data from the Maghreb. A well indurated duricrust indicates favourable climatic conditions for the pedogenic cementation by carbonates of sediment layers at the site after 57 ka. Overlying deposits attributed to the LSA technocomplex yield ages of ~21 ka and ~15 ka, corresponding to the last glacial period, and fall well within the previously established occupation phase in the Maghreb. The last occupation phase at Rhafas took place during the Neolithic and is dated to ~7.8 ka.

In Western Europe, the Middle to Upper Paleolithic transition is associated with the disappearance of Neandertals and the spread of anatomically modern humans (AMHs). Current chronological, behav-ioral, and biological models of this transitional period hinge on the Châtelperronian technocomplex. At the site of the Grotte du Renne, Arcy-sur-Cure, morphological Neandertal specimens are not directly dated but are contextually associated with the Châtelperronian, which contains bone points and beads. The association between Ne-andertals and this " transitional " assemblage has been controversial because of the lack either of a direct hominin radiocarbon date or of molecular confirmation of the Neandertal affiliation. Here we provide further evidence for a Neandertal–Châtelperronian association at the Grotte du Renne through biomolecular and chronological analysis. We identified 28 additional hominin specimens through zooarch-aeology by mass spectrometry (ZooMS) screening of morphologically uninformative bone specimens from Châtelperronian layers at the Grotte du Renne. Next, we obtain an ancient hominin bone pro-teome through liquid chromatography-MS/MS analysis and error-tolerant amino acid sequence analysis. Analysis of this palaeoproteome allows us to provide phylogenetic and physiological information on these ancient hominin specimens. We distinguish Late Pleistocene clades within the genus Homo based on ancient protein evidence through the identification of an archaic-derived amino acid sequence for the collagen type X, alpha-1 (COL10α1) protein. We support this by obtaining ancient mtDNA sequences, which indicate a Neandertal ancestry for these specimens. Direct accelerator mass spectometry radiocarbon dating and Bayesian modeling confirm that the hominin specimens date to the Châtelperronian at the Grotte du Renne. palaeoproteomics | ZooMS | Châtelperronian | Neandertal T o understand the cultural and genetic interaction between the last Neandertals and some of the earliest anatomically modern humans (AMHs) in Europe, we need to resolve the taxo-nomic affiliation of the hominins associated with the " transitional " industries characterizing the replacement period, such as the Châtelperronian (1, 2). The well-characterized Châtelperronian lithic technology has recently been reclassified as fully Upper Pa-leolithic (3) and is associated at several sites with bone awls, bone pendants, and colorants (4, 5). The Grotte du Renne at Arcy-sur-Cure, France, is critical to competing behavioral and chronological models for the Châtelperronian, as at this site the Châtelperronian is stratigraphically associated with hominin remains that are morphologically identified as Neandertals (6–8). Hypotheses explaining this association range from " acculturation " by AMHs (9), to independent development of such artifacts by Neandertals (5), to movement of pendants and bone artifacts from the over-lying Aurignacian into the Châtelperronian layers (10, 11), or to movement of the hominin specimens from the underlying Mousterian into the Châtelperronian layers (10, 12). The first two hypotheses assume that the stratigraphic association of the hominins and the Châtelperronian assemblage is genuine, whereas the latter two hypotheses counter that the association is the result of large-scale, taphonomic movement of material. In all scenarios, the morphological identification of these hominins as Neandertals is accepted but unsupported by molecular evidence. To test the chronostratigraphic coherence of the site, Bayesian models of radiocarbon dates for the site have been constructed (10, 13). The results of these two models contradict each other in the extent to which archaeological material moved between the Châtelperronian and non-Châtelperronian archaeological layers. Furthermore, they have been criticized on various methodological aspects (13, 14), and the first (10) is at odds with some archaeological evidence that suggests that large-scale displacement of material into Significance The displacement of Neandertals by anatomically modern humans (AMHs) 50,000–40,000 y ago in Europe has considerable biological and behavioral implications. The Châtelperronian at the Grotte du Renne (France) takes a central role in models explaining the transition, but the association of hominin fossils at this site with the Châtelperronian is debated. Here we identify additional hominin specimens at the site through proteomic zooarchaeology by mass spectrometry screening and obtain molecular (ancient DNA, ancient proteins) and chronometric data to demonstrate that these represent Neandertals that date to the Châtelperro-nian. The identification of an amino acid sequence specific to a clade within the genus Homo demonstrates the potential of palaeoproteomic analysis in the study of hominin taxonomy in the Late Pleistocene and warrants further exploration.

Objectives: Humans display an 85–95% cross-cultural right-hand bias in skilled tasks, which is considered a derived behavior because such a high frequency is not reported in wild non-human primates. Handedness is generally considered to be an evolutionary byproduct of selection for manual dexterity and augmented visuo-cognitive capabilities within the context of complex stone tool manufacture/use. Testing this hypothesis requires an understanding of when appreciable levels of right dominant behavior entered the fossil record. Because bone remodels in vivo, skeletal asymmetries are thought to reflect greater mechanical loading on the dominant side, but incomplete preservation of external morphology and ambiguities about past loading environments complicate interpretations. We test if internal trabecular bone is capable of providing additional information by analyzing the thumb of Homo sapiens and Pan. Materials and methods: We assess trabecular structure at the distal head and proximal base of paired (left/right) first metacarpals using micro-CT scans of Homo sapiens (n 5 14) and Pan (n 5 9). Throughout each epiphysis we quantify average and local bone volume fraction (BV/TV), degree of anisotropy (DA), and elastic modulus (E) to address bone volume patterning and directional asymmetry. Results: We find a right directional asymmetry in H. sapiens consistent with population-level hand-edness, but also report a left directional asymmetry in Pan that may be the result of postural and/ or locomotor loading. Conclusion: We conclude that trabecular bone is capable of detecting right/left directional asymmetry , but suggest coupling studies of internal structure with analyses of other skeletal elements and cortical bone prior to applications in the fossil record.

Anatomically modern humans replaced Neanderthals in Europe around 40,000 years ago. The demise of the Neanderthals and the nature of the possible relationship with anatomically modern humans has captured our imagination and stimulated research for more than a century now. Recent chronological studies suggest a possible overlap between Neanderthals and anatomically modern humans of more than 5,000 years. Analyses of ancient genome sequences from both groups have shown that they interbred multiple times, including in Europe. A potential place of interbreeding is the notable Palaeolithic site of Riparo Mezzena in Northern Italy. In order to improve our understanding of prehistoric occupation at Mezzena, we analysed the human mandible and several cranial fragments from the site using radiocarbon dating, ancient DNA, ZooMS and isotope analyses. We also performed a more detailed investigation of the lithic assemblage of layer I. Surprisingly we found that the Riparo Mezzena mandible is not from a Neanderthal but belonged to an anatomically modern human. Furthermore, we found no evidence for the presence of Neanderthal remains among 11 of the 13 cranial and post-cranial fragments re-investigated in this study. The process of replacement of Neanderthals by anatomically modern humans around 40,000 years ago in Western Eurasia is one of the most disputed topics in the field of Palaeoanthropology. Although the chronological overlap between the two groups likely lasted more than 5,000 years 1,2 , there is little evidence, if any, of a local coexistence for a significant amount of time. Nonetheless, careful evaluation of geographical scale and of the duration of local interactions between the two populations is critical to assess the possibility of biological admixture and cultural diffusion between 'newly-arrived' moderns and local archaics. As about 2% of Neanderthal ancestry is detected in the genomes of all present-day human populations outside of Africa, the Levant is considered one of the most likely areas where gene flow between Neanderthals and anatomically modern humans could have taken place 3. Additional and regionally confined evidence of hybridization comes from a 37,000–42,000 years old modern human from Peştera cu Oase (Romania), which was shown to have had a Neanderthal ancestor four to six generations earlier based on the analysis of its genome 4. Similar cases of local hybridization have been alleged in various parts of Europe based on anatomical or archaeological evidence, but they have not yet been substantiated by palaeogenetic data 5,6. One example is the late Mousterian site of Riparo Mezzena in northern