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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. sapiens1 or evolved gradually over the last 400 thousand years2. 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.

Evolutionary studies of mammalian teeth have generally concentrated on the adaptive and functional significance of dental features, whereas the role of development on phenotypic generation and as a source of variation has received comparatively little attention. The present study combines an evolutionary biological framework with state-of-the-art imaging techniques to examine the developmental basis of variation of accessory cusps. Scholars have long used the position and relatedness of cusps to other crown structures as a criterion for differentiating between developmentally homologous and homoplastic features, which can be evaluated with greater accuracy at the enamel–dentine junction (EDJ). Following this approach, we collected digital models of the EDJ and outer enamel surface of more than 1000 hominoid teeth to examine whether cusp 5 of the upper molars (UM C5) and cusps 6 and 7 of the lower molars (LM C6 and LM C7) were associated each with a common developmental origin across species. Results revealed that each of these cusps can develop in a variety of ways, in association with different dental tissues (i.e. oral epithelium, enamel matrix) and dental structures (i.e. from different cusps, crests and cingula). Both within and between species variability in cusp origin was highest in UM C5, followed by LM C7, and finally LM C6. The lack of any species-specific patterns suggests that accessory cusps in hominoids are developmentally homoplastic and that they may not be useful for identifying phylogenetic homology. An important and unanticipated finding of this study was the identification of a new taxonomically informative feature at the EDJ of the upper molars, namely the post-paracone tubercle (PPT). We found that the PPT was nearly ubiquitous in H. neanderthalensis and the small sample of Middle Pleistocene African and European humans (MPAE) examined, differing significantly from the low frequencies observed in all other hominoids, including Pleistocene and recent H. sapiens. We emphasize the utility of the EDJ for human evolutionary studies and demonstrate how features that look similar at the external surface may be the product of different developmental patterns. This study also highlights the importance of incorporating both developmental and morphological data into evolutionary studies in order to gain a better understanding of the evolutionary significance of dental and skeletal features.

The mechanism of the Neanderthal extinction and their replacement by modern humans of African origin is one of the most discussed issues in paleoanthropology. Central to this discussion are the questions of the chronological overlap between Neanderthal populations and modern humans in Western Eurasia and the precise geographical circumstances of this overlap. For a long time, the Vindija (Croatia) site was considered to provide solid evidence for a long survival of Neanderthals in Central/Southern Europe. Not only did directly dated Neanderthal remains from layer G1 of the site provide radiocarbon ages postdating the most widely accepted transition time of 40–35,000 radiocarbon years ago (1), but the same layer also yielded a type of split-based bone points commonly assigned to the Aurignacian (2), a stone artefact industry of the early Upper Paleolithic that, to date, only yielded human remains of a modern nature (3). For some, this situation implied the possibility of a long and complex interaction between the two groups of hominins in this region and also falsified the notion of a systematic association between defined archaeological assemblages and specific biological populations at the time of the replacement. In PNAS, Deviese et al. (4) provide new radiocarbon dates for the same Vindija Neanderthal samples, dating them to before 40,000 14C B.P., significantly older than previous efforts dating this material to 29–28,000 and 33–32,000 radiocarbon years (1). The bone points of layer G1 could not be dated, but the range of ages obtained from faunal and human samples in this layer suggests taphonomic mixing as a likely mechanism to explain their stratigraphic association in this part of the Vindija stratigraphic sequence. The situation in Vindija is therefore not at all exceptional, and previous results can be explained by the effect of sample contamination and layer admixture. The first message delivered … [↵][1]1Email: hublin{at}eva.mpg.de. [1]: #xref-corresp-1-1

$\textbf{Objectives:}$ Internal bone structure, both cortical and trabecular bone, remodels in response to loading and may provide important information regarding behavior. The foot is well suited to analysis of internal bone structure because it experiences the initial substrate reaction forces, due to its proximity to the substrate. Moreover, as humans and apes differ in loading of the foot, this region is relevant to questions concerning arboreal locomotion and bipedality in the hominoid fossil record. $\textbf{Materials and methods:}$ We apply a whole-bone/epiphysis approach to analyze trabecular and cortical bone in the distal tibia and talus of $\textit{Pan troglodytes}$ and $\textit{Homo sapiens}$. We quantify bone volume fraction (BV/TV), degree of anisotropy (DA), trabecular thickness (Tb.Th), bone surface to volume ratio (BS/BV), and cortical thickness and investigate the distribution of BV/TV and cortical thickness throughout the bone/epiphysis. $\textbf{Results:}$ We find that $\textit{Pan}$ has a greater BV/TV, a lower BS/BV and thicker cortices than $\textit{Homo}$ in both the talus and distal tibia. The trabecular structure of the talus is more divergent than the tibia, having thicker, less uniformly aligned trabeculae in $\textit{Pan}$ compared to $\textit{Homo}$. Differences in dorsiflexion at the talocrural joint and in degree of mobility at the talonavicular joint are reflected in the distribution of cortical and trabecular bone. $\textbf{Discussion:}$ Overall, quantified trabecular parameters represent overall differences in bone strength between the two species, however, DA may be directly related to joint loading. Cortical and trabecular bone distributions correlate with habitual joint positions adopted by each species, and thus have potential for interpreting joint position in fossil hominoids.

This study explores the morphological differences between the enamel–dentine junction (EDJ) of maxillary and mandibular molars of Neanderthals (n = 150) and recent modern humans (n = 106), and between an earlier Neanderthal sample (consisting of Pre-Eemian and Eemian Neanderthals dating to before 115 ka) and a later Neanderthal sample (consisting of Post-Eemian Neanderthals dating to after 115 ka). The EDJ was visualised by segmenting microtomographic scans of each molar. A geometric morphometric methodology compared the positioning of the dentine horns, the shape of the marginal ridge between the dentine horns, and the shape of the cervix. We also examined the manifestation of non-metric traits at the EDJ including the crista obliqua, cusp 5, and post-paracone tubercle. Furthermore, we report on additional morphological features including centrally placed dentine horn tips and twinned dentine horns. Our results indicate that EDJ morphology can discriminate with a high degree of reliability between Neanderthals and recent modern humans at every molar position, and discriminate between the earlier and the later Neanderthal samples at every molar position, except for the M3 in shape space. The cervix in isolation can also discriminate between Neanderthals and recent modern humans, except at the M3 in form space, and is effective at discriminating between the earlier and the later Neanderthal samples, except at the M2/M2 in form space. In addition to demonstrating the taxonomic valence of the EDJ, our analysis reveals unique manifestations of dental traits in Neanderthals and expanded levels of trait variation that have implications for trait definitions and scoring.

International audience; Terrestrial microvertebrates found in archaeological and palaeontological contexts are well known to be good indicators of palaeoenvironments and climate changes over time. Recent excavations performed at Rhafas Cave in 2008 and 2009 allowed the recovering of microvertebrate remains dating to the Late Pleistocene and Holocene. The main objective of the present work was to evaluate the "microfaunal potential" of the site. Globally, the bone concentration is relatively low, but the diversity is interesting. A preliminary study of selected samples led to the identication of around 764 remains (MNI=147) belonging to more than 20 species of rodents, shrews, elephant-shrews, bats, amphibians, lizards and snakes. The global palaeoecologicalsignal is semi-arid with the maintenance of water ponds in the vicinity of the cave. If the majority of the identied species still inhabit eastern Morocco today, some others are rare or do not occur anymore in the area, leading to interesting palaeoenvironmental and palaeobiogeographic inferences concerning both faunal and human populations.

During the Middle Paleolithic period, carnivores and hominids periodically occupied the same areas at different times and each predator generated significant palimpsests, rendering difficult their archaeological interpretation. Teixoneres Cave, a carnivore den site, located in the northeastern part of the Iberian Peninsula, demonstrates that it is possible to overcome these problems by using a careful strategy in selecting samples for radiocarbon dating, in order to produce an accurate chronology of the site in question and certainly attest the human occupation.

The dispersal of the genus Homo out of Africa approximately 1.8 million years ago (Ma) has been understood within the context of changes in diet, behavior, and bipedal locomotor efficiency. While various morphological characteristics of the knee and ankle joints are considered part of a suite of traits indicative of, and functionally related to, habitual bipedal walking, the timing and phylogenetic details of these morphological changes remain unclear. To evaluate the timing of knee and ankle joint evolution, we apply geometric morphometric methods to three-dimensional digital models of the proximal and distal tibiae of fossil hominins, Holocene Homo sapiens, and extant great apes. Two sets of landmarks and curve semilandmarks were defined on each specimen. Because some fossils were incomplete, digital reconstructions were carried out independently to estimate missing landmarks and semilandmarks. Group shape variation was evaluated through shape–and form-space principal component analysis and fossil specimens were projected to assess variation in the morphological space computed from the extant comparative sample. We show that a derived proximal tibia (knee) similar to that seen in living H. sapiens evolved with early Homo at ∼2 Ma. In contrast, derived characteristics in the distal tibia appear later, probably with the arrival of Homo erectus. These results suggest a dissociation of the morphologies of the proximal and distal tibia, perhaps indicative of divergent functional demands and, consequently, selective pressures at these joints. It appears that longer distance dispersals that delivered the Dmanisi hominins to Georgia by 1.8 Ma and H. erectus to east–southeast Asia by 1.6 Ma were facilitated by the evolution of a morphologically derived knee complex comparable to that of recent humans and an ankle that was morphologically primitive. This research sets the foundation for additional paleontological, developmental, and functional research to better understand the mechanisms underlying the evolution of bipedalism.

We report the application of a molecular barcode method (ZooMS) to identify fragmentary bone remains (>2.5 cm) from a Middle to Upper Palaeolithic sequence at Les Cottes, France. ZooMS uses peptide mass fingerprinting of collagen (the most abundant protein in bone) to discriminate fauna (typically to genus level). Using previously reported peptide markers we initially conducted a blind test using 34 morphologically identified bones, followed by the application of ZooMS on 145 morphologically unidentified bone specimens. For the blind test, ZooMS was in agreement with morphological identifications in all cases, but in some instances taxonomic resolution is lower than morphological identifications. Further, 93.8% (136/145) of spectra obtained for morphologically unidentified bone specimens result in identifications that cannot be taxonomically improved by ZooMS. These include ten bone specimens showing signs of carnivore digestion. Focussing on the unidentified bone specimens of the Châtelperronian unit at Les Cottes (US06), ZooMS identified an additional ≈30% of the total number of bones discovered, increasing the total number of identified bone specimens to 61.8%. Further, ZooMS revealed higher taxonomic richness compared to morphological identifications for US06, thereby providing a more informed interpretation of the faunal community present at Les Cottes during the Châtelperronian.

The study of stone artifacts can provide crucial insights into various aspects of past human adaptation. Knowledge of the geographic provenance of the raw materials, in particular, can inform us of the mobility and resource exploitation strategies employed by past human groups. However, it is difficult to reliably infer dynamic human behaviours from the frequencies with which different raw material are represented at archaeological sites, particularly when the latter reflect the activities of individuals belonging to an extinct human species whose cognitive and physical abilities may have been markedly different from our own.This dissertation provides a new approach for understanding aspects of human adaptation in the Middle Palaeolithic on the basis of toolstone provenance. The approach centers on computer simulations and it is designed to maximize the potential of toolstone provenance information while minimizing reliance on possibly unfounded assumptions about human behaviour derived from studies of present-day societies. The approach is applied to the French Middle Palaeolithic site of the Bau de l’Aubesier, revealing, amongst other things, that the Neanderthal occupants of the site possessed excellent spatial memory and navigational abilities, regularly exploited a surprisingly large area, and appear to have treated stone resources in a strictly utilitarian manner.

The study of stone artifacts can provide crucial insights into various aspects of past human adaptation. Knowledge of the geographic provenance of the raw materials, in particular, can inform us of the mobility and resource exploitation strategies employed by past human groups. However, it is difficult to reliably infer dynamic human behaviours from the frequencies with which different raw material are represented at archaeological sites, particularly when the latter reflect the activities of individuals belonging to an extinct human species whose cognitive and physical abilities may have been markedly different from our own.This dissertation provides a new approach for understanding aspects of human adaptation in the Middle Palaeolithic on the basis of toolstone provenance. The approach centers on computer simulations and it is designed to maximize the potential of toolstone provenance information while minimizing reliance on possibly unfounded assumptions about human behaviour derived from studies of present-day societies. The approach is applied to the French Middle Palaeolithic site of the Bau de l’Aubesier, revealing, amongst other things, that the Neanderthal occupants of the site possessed excellent spatial memory and navigational abilities, regularly exploited a surprisingly large area, and appear to have treated stone resources in a strictly utilitarian manner.

The Iberian Peninsula has been central to the discussion as it was considered to be a 'last refuge' for the species at a time when H. sapiens occupation spread throughout Europe. Much speculation has centred around the idea that extreme climate fluctuations during Marine Isotope Stage (MIS) 3 like Heinrich Stadial 4 (HS-4) were possible drivers for their disappearance. In other words, the extreme warm / cold fluctuations created an uninhabitable setting for Neanderthals, helping the dispersal of H. Sapiens intoIberia. These arguments were originally based on marine and ice core records of far-reaching origin that could provide little information at the local level. At the time this evidence was sufficient, as no other records were available. In recent years, studies have presented terrestrial palaeoclimate records from Iberia during the last glacial, but more are required to adequately understand the local response to Atlantic marine influences. Moreover, few terrestrial records are connected to known archaeological sites. The research presented in this thesis provides a more comprehensive understanding of palaeoclimate change during the Last Glacial Period and the potential implications for Neanderthal / H. sapiens movements in Iberian Peninsula. To achieve this, we examine the importance of a robust and direct U-Th chronology in conjunction with establishing local, terrestrial palaeoclimate records for the coastal Iberian Peninsula during the Holocene and Last Glacial Period.

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 (2007–2017) 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 2007–2014 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.

The central question of this thesis is: What drives late Middle Paleolithic stone artifact variability? In its attempt to answer this question, this thesis is a contribution to understanding variability within and between late Middle Paleolithic assemblages of the European Plain. The selected research area is rich in late Middle Paleolithic open-air sites but up to now lacking detailed quantitative comparisons between them. This thesis uses standardized, more quantitative methods to analyze stone tool variability across techno-complexes. Stone artifact assemblages where analyzed based on a detailed attribute analysis and compared using multivariate methods, like Principal Component Analysis and Nonmetric Multidimensional Scaling. Additionally, methods from 3D geometric morphometrics where applied for the analysis of some individual tool types. The central finding of the studies presented in this thesis is that most of the variability observed in the record is not related to typo-technological classification, and it is not necessarily site-specific nor geographical. The variability can be related to raw materials, but it can also be related to the varying application of similar methods in blank and tool production. The analyses of individual tools suggest further that certain tool concepts, like the bifacial Keilmesser are not restricted to bifacial tools only.

The stable isotope ratios of carbon (d13C), nitrogen (d15N) and sulphur (d34S) are measured to examine human diet, social stratification, mobility and animal husbandry practices. Three locations comprising four sites from the Yellow River Valleys of north China are investigated: Nancheng (Hebei Province), Xishan (Gansu Province), Liyi and Shanren (Shaanxi Province), and this work represents one of the largest and most detailed isotopic research projects ever conducted in China. This thesis focused on time periods and cultures that were previously underrepresented, in the literature such as the early Bronze/Iron Ages to the Qin Dynasty (221-206 BC). Further, this thesis found new evidence concerning dietary patterns, social stratification (or lack thereof), animal husbandry practices and human mobility during these formative pre-Qin Empire periods.

International audience; 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, therefore the emergence of modern human biology is commonly placed at around 200 thousand years ago. The earliest Middle Stone Age assemblages come from eastern and southern Africa but date much earlier. 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. 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 assemblages and almost double the previous age estimates for the lower part of the deposits. 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 Still Bay is a cultural-historic phase within the southern African Middle Stone Age, which remains critical to our understanding of modern human behavioural evolution, as well as to our understanding of the biogeographic spread of different early modern human groups across the southern African sub-continent. However this phase is recognised in the archaeological record solely through the documentation of bifacial stone points (artefacts) that have a specific and complex overall morphology. Variability within this phase, and the accuracy with which the limits of it can be identified in time and in space, has received relatively little to no attention. The purpose of my thesis was to develop an objective and reliable method for identifying and interpreting morphological variability in bifacial points from the key Still Bay assemblages, and further, to test patterns documented with this method against conventional understandings of the phase. The broader central findings suggest that Still Bay point morphology varies significantly but uniformly through time, and discretely across space between bifacial point regional groupings. Therefore, treatment of the Still Bay as a culturally homogeneous entity and marker of demographic affiliation may be unreliable in a broader regional context.

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, behavioral, 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 Neandertals 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 zooarchaeology 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 proteome 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.

Diets and food are indisputably core facets of human society. The great apes still rely on plants to supply most of their nutritional needs. Humans, however consume a diet that is nearly unrecognizable from that of early hominin and human ancestors. While the virtues of plant foods are widely extolled, it is done so without a regard for how wild and unrefined plant foods must have contributed to the diets of our pre-agricultural ancestors. The purpose of my dissertation was to examine how wild African tubers, which are targeted by the Hadza hunter-gatherers of Tanzania, contribute the nutritional elements that are essential for health and survival in the East African mosaic environment. To accomplish this objective, I conducted three projects. First, I tested the efficacy of brief roasting, used by the Hadza, on starch gelatinization. Second, I submitted four species of Hadza tubers to an in-vitro simulation of the gastro-intestinal tract, and measured the bioaccessibility of glucose from digestion. Finally, I led an analysis of the Hadza gut microbiome to explore the polysaccharide degrading potential of mutualistic microbiota and learn whether Hadza may be better provisioned by metabolites from fermentation of the residual indigestible plant fibers in their diet.

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.

Prehistoric dental treatments have been known from the Neolithic 9,000-7,500 years before present (BP) [1], when the adoption of early farming culture caused an increase of carious lesions [2]. Only a few early cases have been documented [3], some were characterized by in vivo perforation of the crown surface made by a drilling tool [1]. Here we document the earliest evidence of proto-dental therapeutic intervention on a Late Upper Paleolithic modern human lower right third molar (RM3) from a burial in Northern Italy [4]. The RM3 belongs to a young male individual (ca. 25 years old) unearthed in 1988 from the Epigravettian deposit of Riparo Villabruna (Sovramonte – Belluno, Italy), and dated around 14,160-13,820 BP. This tooth presents a large occlusal cavity, with a polished internal surface and extensive enamel chipping traces on the steep mesial wall. Within the cavity four caries are present. The cavity is sub-squared on the lingual and mesial sides but rounded on the buccal and distal sides. Using Scanning Electron Microscopy (SEM) we show the presence of striations within the cavity, which fade out towards the occlusal surface probably as a consequence of tooth wear. The striations have a ”V” shaped transverse section and microstriations at the bottom, sharply defined, with a high apex, steep sides, narrow cross-sections and well-defined parallel ancillary ridging, as typically displayed by cutmarks on teeth [5]. Based on in vitro experimental replication and a complete functional reconstruction of the Villabruna dental arches, we confirm that the identified striations and the associated extensive enamel chipping on the mesial wall of the cavity were produced ante-mortem by pointed flint tools during scratching and chiseling activities. The Villabruna specimen is therefore the oldest known evidence of dental caries intervention, suggesting rudimentary knowledge of disease treatment well before the Neolithic. This study also suggests that primitive forms of carious treatment in human evolution entail an adaptation of the well-known toothpickings for levering and scratching rather than drilling practices.