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International audience; This article proposes a first synthesis of ancient funerary practices in western Normandy based on a corpus of 29 sites selected from a very dense documentation of uneven quality. Most of them are unpublished, some have been partially published. This corpus illustrates a wide range of funerary practices identified at regional level and covers a vast chronological range from the 1st century BC to the 5th century AC.The cremation of corpses largely dominates the Lower Normandy funerary landscape since the Conquest and throughout the Early Roman Empire. This practice gradually gave way to that of inhumations, which in turn became predominant in the 2nd century. However, numerous sites prove that these two rites coexisted for a long time. Beyond these considerations, the ancient funerary sites of western Normandy reflect a variety of occupations from the isolated tomb to the family mausoleum, via the community necropolis. Only the urban necropolises remain undocumented to this day.; Le dernier bilan de la recherche archéologique portant sur le fait funéraire antique en Normandie a été mené entre 2004 et 2010. Il a mis en évidence un fort potentiel d'étude avec pas moins de 41 indices. En quelques années, ce chiffre a triplé du fait de l'essor de l'archéologie préventive sur le territoire normand. Fin 2015, ce sont pas moins de 158 sites funéraires recensés dont 50 % des découvertes, soit près de 80 sites, sont postérieures à 1984. Parallèlement, peu de publications ont vu le jour ces trois dernières décennies. L'atelier "funéraire" du PCR Arbano s'est donné comme objectif de prendre en compte la totalité des découvertes anciennes (études complémentaires) comme récentes (sites inédits) et de dresser un premier état des connaissances sur le fait funéraire antique en Normandie occidentale.

Genomic analyses of Neanderthals have previously provided insights into their population history and relationship to modern humans1–8, but the social organization of Neanderthal communities remains poorly understood. Here we present genetic data for 13 Neanderthals from two Middle Palaeolithic sites in the Altai Mountains of southern Siberia: 11 from Chagyrskaya Cave9,10 and 2 from Okladnikov Cave11—making this one of the largest genetic studies of a Neanderthal population to date. We used hybridization capture to obtain genome-wide nuclear data, as well as mitochondrial and Y-chromosome sequences. Some Chagyrskaya individuals were closely related, including a father–daughter pair and a pair of second-degree relatives, indicating that at least some of the individuals lived at the same time. Up to one-third of these individuals’ genomes had long segments of homozygosity, suggesting that the Chagyrskaya Neanderthals were part of a small community. In addition, the Y-chromosome diversity is an order of magnitude lower than the mitochondrial diversity, a pattern that we found is best explained by female migration between communities. Thus, the genetic data presented here provide a detailed documentation of the social organization of an isolated Neanderthal community at the easternmost extent of their known range.

International audience; The oral microbiota is a highly complex and diversified part of the human microbiome. Being located at the interface between the human body and the exterior environment, this microbiota can deepen our understanding of the environmental impacts on the global status of human health. This research topic has been well addressed in Westernized populations, but these populations only represent a fraction of human diversity. Papua New Guinea hosts very diverse environments and one of the most unique human biological diversities worldwide. In this work we performed the first known characterization of the oral microbiome in 85 Papua New Guinean individuals living in different environments, using a qualitative and quantitative approach. We found a significant geographical structure of the Papua New Guineans oral microbiome, especially in the groups most isolated from urban spaces. In comparison to other global populations, two bacterial genera related to iron absorption were significantly more abundant in Papua New Guineans and Aboriginal Australians, which suggests a shared oral microbiome signature. Further studies will be needed to confirm and explore this possible regional-specific oral microbiome profile.

International audience; The WomenSOFar ANR project (Individual life histories and WOMEN Status at the Onset of FARming. Bioarchaeological perspectives in the French and Mediterranean Prehistoric context) proposes to discuss the place of women in first farmers’ communities by using a multidisciplinary approach. It aims at deciphering how individuals move, eat, care for and work and combines previous and new dataset on bioanthropology, biogeochemistry, bioarchaeology and archaeology. The project focuses on specific questions as: 1- Does maternal/allomaternal investment and food transfers differ according to their own experience and local subsistence economy? 2- Does parental/alloparental investment lead to female vs. male social distinction as early as childhood? Does a timing/pattern exist to get the role of “female” or “male” into these communities? 3- Why do women migrate differently from men and how does it influences their diet and its diversity? 4- Are women involved in specific activities affecting their health and morphology? Does this exist in connection with their place of origin? Does it have an impact on their diet/the emergence of their childhood socialization? The project investigates human remains and their cultural contexts from several sites located in different natural environments in France and North Africa, mainly dated to the 5th millennium BCE. Skeletal and dental records, cross-sectional geometry of long bones, multi-element isotope ratios, funerary records, dental calculus micro-remains and microbiome and kinship data will be compared. Examples of previous researches using this multidisciplinary approach illustrates the risk/gain of such study; among other things, data allowed to identify female individuals with specific life history and mobility pattern and male individuals with possible “disadvantaged” status. WomenSOFar expects to provide a new vision of the diversity and mutability of women status in Prehistoric times at the onset of major societal disruptions.

International audience; ObjectivesThe shape of the trapezium and first metacarpal (Mc1) markedly influence thumb mobility, strength, and the manual abilities of extant hominids. Previous research has typically focused solely on trapezium-Mc1 joint shape. Here we investigate how morphological integration and shape covariation between the entire trapezium (articular and non-articular surfaces) and the entire Mc1 reflect known differences in thumb use in extant hominids.Materials and MethodsWe analyzed shape covariation in associated trapezia and Mc1s across a large, diverse sample of Homo sapiens (n = 40 individuals) and other extant hominids (Pan troglodytes, n = 16; Pan paniscus, n = 13; Gorilla gorilla gorilla, n = 27; Gorilla beringei, n = 6; Pongo pygmaeus, n = 14; Pongo abelii, n = 9) using a 3D geometric morphometric approach. We tested for interspecific significant differences in degree of morphological integration and patterns of shape covariation between the entire trapezium and Mc1, as well as within the trapezium-Mc1 joint specifically.ResultsSignificant morphological integration was only found in the trapezium-Mc1 joint of H. sapiens and G. g. gorilla. Each genus showed a specific pattern of shape covariation between the entire trapezium and Mc1 that was consistent with different intercarpal and carpometacarpal joint postures.DiscussionOur results are consistent with known differences in habitual thumb use, including a more abducted thumb during forceful precision grips in H. sapiens and a more adducted thumb in other hominids used for diverse grips. These results will help to infer thumb use in fossil hominins.

International audience; Présentation des résultats préliminaires de travail de terrain effectué dans la région du Tohoku au Japon d'octobre 2022 à mars 2023.

Boves Castle, located in northern France, was occupied between the 9th and 16th centuries AD. Under excavation since 1996, this high-status site has yielded more than six tons of faunal remains. Until now, archaeozoological studies have focused on the early occupation of the site (10th and 11th centuries AD), revealing specific supply and consumption strategies, widely based around birds. Aside from the very large amount of domestic fowl, Boves Castle is distinguished by the number and diversity of wild birds from various environments, including wetland, coastal, forested and anthropized areas. However, the special feature of this site is the importance of freshwater birds, especially large waders, which remained a status symbol until modern times in this region. On the basis of this outstanding example, and other high-status sites from the same region, we aim to discuss the importance and status of various domestic and game bird categories in northern France during the High Middle Ages. We will also address the subject of controlled hunting areas around castles and other high-status consumer sites, for a better understanding of the link between power and consumption during the medieval period.

Previous research indicates that the human genetic diversity found in Wallacea - islands in present-day Eastern Indonesia and Timor-Leste that were never part of the Sunda or Sahul continental shelves - has been shaped by complex interactions between migrating Austronesian farmers and indigenous hunter-gatherer communities. Here, we provide new insights into this region’s demographic history based on genome-wide data from 16 ancient individuals (2600-250 yrs BP) from islands of the North Moluccas, Sulawesi, and East Nusa Tenggara. While the ancestry of individuals from the northern islands fit earlier views of contact between groups related to the Austronesian expansion and the first colonization of Sahul, the ancestry of individuals from the southern islands revealed additional contributions from Mainland Southeast Asia, which seems to predate the Austronesian admixture in the region. Admixture time estimates for the oldest individuals of Wallacea are closerto archaeological estimates for the Austronesian arrival into the region than are admixture time estimates for present-day groups. The decreasing trend in admixture times exhibited by younger individuals supports a scenario of multiple or continuous admixture involving Papuan- and Asian-related groups. Our results clarify previously debated times of admixture and suggest that the Neolithic dispersals into Island Southeast Asia are associated with the spread of multiple genetic ancestries.

Summary The integration of the growth of the brain and the cranial base suggests that each system may influence the other, notably during the first three years of life, although this influence has never been proven to be exclusive. The aim of our work was to analyse the dynamics of normal growth on the one hand, and the development and ontogenetic allometry of the cranial base in the infant on the other hand. Material and method A total of 32 infants (17 males/15 femeles) having been included in the unexpected infant death french protocol were analyzed. Three-dimensional reconstructions of the cranial base were performed from CT scans. The technique combined manual segmentation of regions of interest, contour extraction and surface reconstruction. Nineteen landmarks were positioned on each of the bone surfaces. Results No correlation was observed between sex assigned at birth and shape, weight, crown-heel length, or head circumference. Principal component analysis showed that 85.5% of the variance observed on the first component was secondary to growth. After Procrustes superimposition, 25% of the shape variance observed was explained by the first principal component. It showed anteroposterior lengthening of the cranial base. In addition, the height, width and length of the posterior fossa increased and the relative position of the basion was displaced inferiorly and anteriorly with flexion of the sphenoid angle. Negative allometry was also observed. Conclusion Our study, carried out in a rigorously selected population of infants, presents a fundamental approach to ontogeny through study of shape, growth and ontogenetic allometry.

International audience; Understanding the contribution of fish to the human dietis an important debate in past population studies as it implies specific organization (e.g. adapted techniques, gender-based activities?) and relationship with the local or sublocal environments (e.g. mobility, seasonal-based exploitation). In the Paris Basin, the role of fish has been discussed mainly from rare but obviousarchaeologicalremains such as fish bones or hooks. Previous isotopic data from Neolithic human bone bulk collagen highlighted the potential of freshwater resources in protein diet but were not able to quantify it or even confirm its role. In this study we conducted, for the first time on early farmers from France,compound specific isotope analysis on bone collagen amino acids (CSIA-AA). Results indicate a particularly high trophic level of some humans with protein intake mainly based on terrestrial animal resources (domestic cattle and pig), while others rather havea plant-based diet. Freshwater resources appeared to be not significant and several possible food combinations are discussed. Pig particularly appears more herbivorous than previously assumed and has possibly consumed manured cereals from human food waste.

International audience; Les examens paléopathologiques de sujets immatures se confrontent au caractère peu spécifique des modifications osseuses observables macroscopiquement. Afin d’aider à la caractérisation des pathologies des sujets immatures, le CRAHAM a mené de 2020 à 2022 le programme de recherche ANCESTOR en collaboration avec la plate-forme d’imagerie CYCERON, le centre de microscopie CMABio3, le laboratoire CRISMAT et en partenariat avec La Fabrique de patrimoines en Normandie. Cette recherche, financée par l’Union européenne dans le cadre du programme opérationnel FEDER/ FSE 2014-2020, avait pour objectif d’obtenir des protocoles fiables d’analyse de pathologies osseuses, de troubles de croissance et de pathologies carentielles des enfants entre 0 et 5 ans. Envisagé pour constituer une aide à une meilleure évaluation paléopathologique des populations normandes de l’Antiquité à la période moderne, ce programme a permis de constituer un référentiel ostéologique s’appuyant sur la complémentarité des examens macroscopiques, radiographiques, microtomographiques et histologiques. Par la mise en place d’un protocole descriptif standardisé, nous avons cherché à identifier le pouvoir prédictif de l’aspect de surface du périoste sur les aspects osseux visibles par radiographie, microtomographie et histologie. Pour cela, nous nous sommes appuyés sur un corpus de 39 individus décédés avant l’âge de 3 ans et inhumés dans des cimetières normands médiévaux et modernes. Les résultats montrent que les descriptions macroscopiques sont faiblement associées aux autres descriptions et que la combinaison de plusieurs méthodes d’analyse est indispensable pour établir un lien entre l’aspect de surface et les modifications tissulaires. Ces résultats permettront d’orienter les choix méthodologiques dans l’optique d’élaborer un modèle interprétatif logiciste de diagnostic utilisable par la communauté archéo-anthropologique pour l’identification de maladies ou de carences osseuses des sujets immatures.

Determining how sibling interactions alter the fitness outcomes of dispersal is pivotal for the understanding of family living, but such studies are currently scarce. Using a large demographic dataset on pre-industrial humans from Finland, we studied dispersal consequences on different indicators of lifetime reproductive success according to sex-specific birth rank (a strong determinant of dispersal in our population). Contrary to the predictions of the leading hypotheses, we found no support for differential fitness benefits of dispersal for either males or females undergoing low vs. high sibling competition. Our results are inconsistent with both hypotheses that family members could have different fitness maximizing strategies depending on birth rank, and that dispersal could be mainly driven by indirect fitness benefits for philopatric family members. Our study stresses the need for studying the relative outcomes of dispersal at the family level in order to understand the evolution of family living and dispersal behaviour.

e report a genome-wide association study of facial features in >6000 Latin Americans based on automatic landmarking of 2D portraits and testing for association with inter-landmark distances. We detected significant associations (P-value

International audience; Humans have evolved to interact smoothly with their environment and with others. There are highly complex processes that enable these interactions and many systems are engaged, from the peripheral somatosensory system to a distributed network of cortical regions. This chapter will address the pathway from the peripheral receptors to the brain, including steps where there is the potential for the processing and integration of information, as well as why these have occurred in our evolution. We have a vast system of somatosensory afferents that are distributed over our skin and in our bodies to capture precise signals about our interactions with the world. Somatosensory afference comes from mechanical, thermal, and chemical stimuli, where numerous different types of somatosensory afferent, namely those that respond to mechanoreceptive, thermoreceptive, and/or nociceptive signals respectively, register specific contact and behavior. These signals are persistent and there is considerable integration of this information even before it reaches the brain. Once the input reaches sub-cortical structures and is passed on to the cerebral cortex, there is again a wealth of processes that interact smoothly, to produce the awareness of our body in space and its interactions. These include principal somatosensory targets, such as the primary (S1) and secondary (S2) somatosensory cortices, and the insula. In the integration of information, many other regions are involved, including the activation of a range of cognitive (e.g. attention, memory, learning) and emotional/affective mechanisms, as well as multisensory processing. This chapter will consider the intricacies of these processes, by exploring the sensory origin of body perception.

International audience; Large-scale disasters such as the 2011 Tohoku Earthquake and Tsunami have produced fatalities in large quantities whose bodies had to be searched for, recovered, identified, and returned to their families. The care and management of these bodies have many implications, including the legal and administrative requirements of accounting for the missing and the dead and the social need to mourn for the dead individually and collectively. For the first aspect, forensics has a central role in applying scientific methods and techniques to recover and identify the dead, enabling, for example, the issuance of a death certificate required by the victim's family to claim succession and insurance money. And for the latter, religion has played a traditional role in organising and prescribing the funerary practices and norms for mourning. Moreover, religion can largely influence how a disaster is experienced and made sense of (fatality, punishment). The present research focuses on the issues that arise from the interaction between religious and cultural funeral practices and the scientific and legal forensic management of the dead. To what extent do the internationally elaborated forensic protocols for handling and caring for the dead effectively adjust to, negotiate with, or oppose local funerary prescriptions?

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International audience; Asymptomatic variations (AV) in bones and teeth are widely used in biological anthropology, particularly to understand intra- and inter-population affinities.This study focuses on the first step of any study: the repeatability and reproducibility of the method implemented. Using the Kappa coefficient, the reliability of 198 variables divided into three anatomical areas (cranial, post-cranial and teeth) was analyzed. The sample used was selected among the best represented and best preserved individuals of the Notre-Dame-du-Bourg osteo-archaeological collection (Digne-les-Bains, medieval and modern period). The analyses were performed after four rounds of scoring (three observers), and when more than 20 observations of polymorphic variations were obtained. The quality of agreement was established according to the Fleiss et al. scale (2003). Although missing data were a hindrance in this study, several proposals were made to overcome the problem. We also highlight and discuss the fact that experience has an impact on the observation of qualitative variables. The outcome is a proposal for a protocol based on selecting repeatable and reproducible AVs to normalize comparisons between studies in both osteoarchaeological and forensic contexts.; Les variations asymptomatiques (VA) osseuses et dentaires sont grandement utilisées en anthropologie biologique, notamment pour appréhender les affinités intra- et inter-populationnelles. Cette étude se concentre sur la première étape de toute étude : la répétabilité et la reproductibilité de la méthode mise en place. À partir de l’utilisation du coefficient de Kappa la fiabilité de 198 variables réparties en trois zones anatomiques (crâne, post-crâne et dents) a été analysée. L’échantillon utilisé a été sélectionné parmi les individus les mieux représentés et conservés de la collection ostéo-archéologique Notre-Dame-du-Bourg (Digne-les-Bains, périodes médiévale et moderne). Les analyses ont été réalisées, suite à quatre séries de cotations (trois observatrices), et lorsque plus de 20 observations sur des variables polymorphiques ont été obtenues. La qualité de l’accord a été établie selon l’échelle de Fleiss et al. (2003). Bien que les données manquantes aient été un frein dans cette étude, plusieurs propositions sont faites afin de palier ce phénomène. De plus, nous mettons en avant et discutons le fait que l’expérience a un impact sur l’observation des variables qualitatives. Au final, un protocole basé sur la sélection de VA répétables et reproductibles est proposé pour normaliser les comparaisons entre études dans un contexte aussi bien bioarchéologique que médico-légal.

International audience; The Bayesian statistical approach considers teeth as forming a developmental module, as opposed to a tooth-by-tooth analysis. This approach has been employed to analyze Upper Pleistocene hominins, including Neandertals and some anatomically modern humans, but never earlier populations. Here we show its application on five hominins from the TD6.2 level of the Gran Dolina site (Homo antecessor, Early Pleistocene) and the Sima de los Huesos site (Middle Pleistocene) of the Sierra de Atapuerca (Burgos, northern Spain). Our results show an advanced development of the third molars in both populations with respect to modern Homo sapiens. In addition, the Sima de los Huesos hominins differ from H. sapiens and H. antecessor in the relatively advanced development of their second molar. The relative mineralization of I1/M1 in H. antecessor appears to be similar to that of modern humans, as opposed to that of Neandertals, which appear to be unique. These observations, combined with reduced enamel formation times and the advanced development of the third molars, appear to indicate a shorter ontogenetic period in the hominins from Gran Dolina and Sima de los Huesos in comparison to modern human average.

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The present article aims to introduce the IATEKA programme “Interdisciplinary Approach to the Territorial Evolution of the Kvarner Archipelago” and its first results, started in 2018 and focused on examining the long-term relationship between humans and their environment, with a particular focus on the construction, management and transformation of the island territories and landscapes of the Kvarner Archipelago, as well as the fluctuation of the different populations and their impact and/or adjustment to the particular environment. As an interdisciplinary programme, methods and results obtained in the field of sedimentary and landscape archaeology, physical anthropology or bioarchaeology, specific analyses such as stable isotopes, ancient DNA, paleobiology and sedimentology are brought to a new level of inter-examination. Although this programme aims to approach the questions of human-environment interactions over a long, or even very long period in as comprehensive a manner as possible and through an interdisciplinary approach, the article focuses only on the results of the palynological studies, relating to the period between Antiquity and the Early Middle Ages. Indeed, the results obtained on the lake sediments allow us to better understand the evolution of the environment of one of the islands of the archipelago, the island of Cres. It also seemed impossible to approach the changes observed for the period of Late Antiquity stricto sensu without addressing, even if only briefly, the character traits of the region that were established from the beginning of Antiquity and the effective presence of Roman power in the region.

International audience; The contribution of shellfish to ancient subsistence has been overlooked in many archaeological studies in Central-East Polynesia. Archaeomalacology, however, can shed light on a wider range of exploited mollusks. In this paper we investigate the exploitation of the chiton Acanthopleura gemmata from the Hane Dune site, Ua Huka, Marquesas Islands. Although several previous studies have highlighted the importance of chitons in Hane’s archaeological record, little information has been presented about this important subsistence item as it is seldom identified with precision and is usually quantified by weight. Using zooarchaeological methods, spatial analysis, and biometry, we demonstrate the importance of chitons in past ‘Enata subsistence. We highlight the intensive exploitation of A. gemmata during the first three centuries of occupation of the site, beginning ca. AD 950, until its abrupt depletion, correlated with the progressive exploitation of the limpet Cellana radiata. Through this case study, we stress the importance of precise taxonomic identification and refined recovery protocols to better understand subsistence trends.

International audience; Archaeological excavations carried out in the 1990s in the former rural parish of Rigny (Indre-et-Loire, France) uncovered 1738 graves. Among the individuals discovered, a female adult presents a series of cranial deformities characteristic of an extremely rare pathology, til then unknown in the archaeological record: the Treacher-Collins syndrome. It induces facial dysmorphism and is generally associated with hearing loss or even conductive deafness of the patient due to malformations of the hearing system.The retrospectif diagnosis of this individual was based on macroscopic analysis and computed tomography (CTscan). The use of 3D imaging made it possible to explore the impact of the pathology on the individual's hearing.In addition to the functional implications, the negative impacts of facial dysmorphic disorders in the social sphere are well known in today's societies while they are poorly documented for the medieval period.During this period, diet was an important social marker. The analysis of the individual affected by this syndrome’s diet, compared to those of other members of the Rigny village community, provide an opportunity to discuss the social significance of disability in a rural context in the Middle Ages. In this perspective, the isotopic ratios of carbon and nitrogen in this individual's bone collagen were compared to those of forty-seven other individuals found on the site.Archaeological, paleopathological and biochemical information are relevant to better characterize the functional repercussions of this syndrome on the Rigny individual and also to discuss the integration of this individual into this medieval village community.

International audience; Within the framework of the WP2 of the A*Midex TRIADS programme, supported by the National Heritage Institute of Tunisia, the University of Sfax, the University of Sousse and Aix-Marseille University, a series of four kakemonos presenting the main results of the excavation and study of funerary urns conserved at the Archaeological Museum of Sfax and coming from the archaeological site of Thaenae has been made for temporary and itinerant exhibitions aimed at the general public. These posters were presented for the first time during the closing day of the TRIADS programme on 17 February 2023 at the Institut National du Patrimoine de Tunisie.; Dans le cadre du WP2 du programme A*Midex TRIADS, porté par l’Institut National du Patrimoine, l’Université de Sfax, l’Université de Sousse et Aix-Marseille Université, une série de quatre kakemonos présentant les principaux résultats de la fouille et de l’étude des urnes funéraires conservées au musée archéologique de Sfax et provenant du site archéologique de Thaenae a été confectionnée pour des expositions temporaires et itinérantes à destination du grand public. Ces panneaux ont été présentées pour la première fois lors de la journée de clôture du programme TRIADS le 17 février 2023 à l’Institut National du Patrimoine de Tunisie.

International audience; The estimation of a biological profile in biological anthropology is a central point inthe study of human remains. To specify this profile, the observation of variations, and patholo-gies on the skeleton is a tool that makes an individual unique. In this research, the focuswas on the distal part of the humerus. Through the study of five individuals from an osteo-archaeological collection, conformational anomalies of the medial epicondyle of the humeruswere highlighted. The aim is to describe them and to propose an etiology of appearance. Forthis purpose, observation of the rest of the skeleton was carried out, in order to search forthe presence of other variations, in particular anomalies of fusion of secondary ossificationcenters. Overall, a non-fusion of the medial epicondyle of the humerus, two fused medialmicro-epicondyles and two supplementary fusions were observed. These are very rare anoma-lies, with an estimated prevalence of 0.7%. Conformational abnormalities of this area can leadto pain. Therefore, medical imaging for diagnostic purposes may be available: useful data forantemortem documentation in case of a disturbing disappearance. Following the study of theentire skeleton, a common etiology for these different conformations cannot be proposed. How-ever, two categories of etiologies can be proposed: a congenital anomaly and a developmentalanomaly with an environmental influence (mechanical or deficiency). The new knowledge ofthis area will enrich the use of individualization factors, especially in the context of forensicanthropology.

Réflexion sur l'approche par compétences : une formation par la (recherche)-création (2022-2023).

Leprosy, a chronic infectious disease, leads to blood mineral imbalances: low levels of zinc, calcium, magnesium, and iron and high levels of copper. Interestingly, in late medieval Europe, minerals were used to treat leprosy. We investigated physiological responses to leprosy and possible evidence of treatment in dental tissues of leprosy sufferers from medieval Denmark and early 20th century Romania when multidrug therapy was not then yet invented. Using Synchrotron Fluorescence (SXRF) and laser ablation (LA-ICP-TOFMS), we show marked covariations in the zinc, calcium, and magnesium distributions, which are compatible with clinical studies but cannot be directly attributed to leprosy. Minerals used historically as a treatment for leprosy show no detectable intake (arsenic, mercury) or a diffuse distribution (lead) related to the daily consumption of contaminated water and food. Intense lead enrichments indicate acute incorporations, potentially through the administration of lead-enriched medication or the mobilization of lead from bone stores to the bloodstream during intense physiological stress related to leprosy. However, comparisons with a healthy control group are needed to ascertain these interpretations. The positive correlations and the patterns observed between lead and essential elements may indicate underlying pathophysiological conditions, demonstrating the potential of the two techniques for investigating diseases in past populations.

International audience; Chez les primates, les adaptations locomotrices sont nombreuses et autorisent l’accès à des milieux variés et complexes. La morphologie externe du squelette et la structure interne des os longs reflètent ces adaptations, liées à certains comportements locomoteurs. L’étude de la covariation de ces traits est susceptible d’expliquer des mécanismes évolutionnaires à l’origine de ces phénotypes. Une forte covariation peut être vue comme une modalité d’adaptation à une fonction ou à un environnement sur le temps long ; tandis qu’une faible covariation peut être interprétée comme un indice de plasticité phénotypique. L’objectif de cette étude est de déterminer des schémas d’adaptation au sein des catarrhiniens par l’étude de la covariation entre la morphologie externe et la structure interne du système fémoro-pelvien. L’étude porte sur le complexe fémoro-pelvien de 79 primates catarhiniens adultes représentant 6 genres (Homo, Pan, Gorilla, Hylobates, Papio et Macaca). Nous avons analysé les niveaux de covariation entre des paramètres structurels de la diaphyse fémorale (paramètres de sections diaphysaires) et les morphologies fémorale et pelvienne. Principaux résultats : (1) les taxons aux répertoires proches ne présentent pas nécessairement un patron de covariation morpho-structurelle comparable. (2) la covariation morpho-structurelle n’est pas nécessairement corrélée au niveau de spécialisation locomotrice. Les catarrhiniens présentent une grande variété de patrons de covariation morpho structurelle au sein du système fémoro pelvien. Seuls les taxons pratiquant une locomotion quadrupède terrestre associée à des comportements arboricoles présentent une covariation entre la morphologie du pelvis et la distribution de l’os cortical dans la diaphyse fémorale. Chez les humains, la forte présence de covariation morpho-structurelle au niveau du fémur serait elle une particularité du mode bipède ?

International audience; The Cultural Transmission of Reproductive Success (CTRS) is one of the various cultural processes that can impact human genetic evolution. In this process, individuals from large families have more children on average. Here, we develop and evaluate methods to infer this process from genomic data, using two approaches: (1) Approximate Bayesian computation, which uses summary statistics computed on inferred genealogies from genomic data and (2) deep neural networks, which are directly trained on genomic data. These methods rely on large simulated datasets incorporating varying levels of CTRS. Both competing approaches show a good ability to infer CTRS on genomic data and worth investigating under more complex evolutionary histories.

International audience; While the exploitation of aquatic fauna and flora has been documented in several primate species to date, the evolutionary contexts and mechanisms behind the emergence of this behavior in both human and non-human primates remain largely overlooked. Yet, this issue is particularly important for our understanding of human evolution, as hominins represent not only the primate group with the highest degree of adaptedness to aquatic environments, but also the only group in which true coastal and maritime adaptations have evolved. As such, in the present study we review the available literature on primate foraging strategies related to the exploitation of aquatic resources and their putative associated cognitive operations. We propose that aquatic resource consumption in extant primates can be interpreted as a highly site-specific behavioral expression of a generic adaptive foraging decision-making process, emerging in sites at which the local cost-benefit trade-offs contextually favor aquatic over terrestrial foods. Within this framework, we discuss the potential impacts that the unique intensification of this behavior in hominins may have had on the evolution of the human brain and spatial ecology.

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International audience; Scientific examination of the heart of Blessed Pauline Jaricot—a French missionary figure—was carried out in 2022. As tandem mass spectrometry proteotyping has proven to be valuable to obtain the broad taxonomic repertoire of a given sample without any a priori information, we aimed at exploring the conditions of preservation of the relics and possible conditions of death. Metaproteomics and high-resolution microtomography imaging approaches were combined. A dataset comprising 6731 high-resolution MS/MS spectra was acquired and 968 of these spectra could be assigned to specific peptidic biomolecules. Based on the taxonomical information encompassed by the identified peptide sequences, 5 phyla were identified amongst eukaryota (94% of the biomass): Ascomycota (55%), with the species Aspergillus versicolor, Trichophyton mentagrophytes and Aspergillus glaucus, corresponding to expected cadaverous fungal flora; Chordata (42%), represented by a unique species, Homo sapiens; Streptophyta (3%); and Arthropoda (traces). Bacteria (6% of the biomass) were poorly represented. No trace of embalming substance could be retrieved, nor any pathogens. Imaging evidenced no heart defect nor embalming traces. No evidence that was inconsistent with natural and spontaneous conservation could be retrieved. This study prefigures the power of modern molecular techniques such as paleoproteotyping coupled to microtomography to gain insight into historical relics.

The Longobard necropolis of Castel Trosino dates from the 6th to the 8th century CE. Among the tombs excavated, the skull of an older female shows the first evidence of a cross-shaped bone modification on a living subject. Macroscopic, microscopic, and CT scan analyses revealed signs of at least two sets of scraping marks. Specifically, SEM analysis shows that perimortem bone-scraping traces are present on the skull. Both healed and non-healed defects suggest that the woman has received at least twice intentional bone modifications to address her condition. This is the first evidence of a cross-shaped therapeutic intervention on a living subject.

International audience; La compilation des dates 14C disponibles pour les différents types de sépultures collectives du bassin nord occidental de la Méditerranée a conduit à l’élaboration d’une première base de données de 103 sites archéologiques, 117 contextes funéraires et 199 dates 14C. Leur analyse critique nous a finalement conduits à poursuivre l’étude sur un ensemble de 83 sites archéologiques (80,6 %), 91 sépultures collectives (77,8 %) et 166 dates 14C (83,4 %). La typologie établie se compose de six grandes catégories de sépultures collectives : dolmen, grotte, grotte aménagée, hypogée, fosse simple et fosse complexe. Bien que le nombre de sépultures datées par le radiocarbone soit limité par rapport au nombre total de sites funéraires, quelques tendances semblent se dégager. Les dolmens et les grottes aménagées ou non sont utilisés de façon synchrone dès la seconde moitié du 4e millénaire. C’est également le cas des fosses complexes mais celles-ci ne sont documentées qu’en Catalogne. Les hypogées et les fosses simples semblent apparaître un peu plus tardivement. Cette synthèse n’est qu’une étape de travail qui permettra de circonscrire les orientations à donner à cette thématique, notamment par une multiplication des datations radiocarbones (particulièrement des dolmens catalans) et la prise en compte des données chrono-culturelles lorsqu’elles sont disponibles.

It has been shown that living in risky environments, as well as having a risky occupation, can moderate risk-tolerance. Despite the involvement of dopamine in the expectation of reward described by neurobiologists, a GWAS study was not able to demonstrate a genetic contribution of genes involved in the dopaminergic pathway in risk attitudes and gene candidate studies gave contrasting results. We test the possibility that a genetic effect of the DRD4-7R allele in risk-taking behavior could be modulated by environmental factors. We show that the increase in risk-tolerance due to the 7R allele is independent of the environmental risk in two populations in Northern Senegal, one of which is exposed to a very high risk due to dangerous fishing.

The WomenSOFar ANR project (2022-2026; ANR-21-CE03- 0008; Individual life histories and WOMEN Status at the Onset of FARming. Bioarchaeological perspectives in the French and Mediterranean Prehistoric context) proposes to discuss the place of women in first farmers’ communities by using a multidisciplinary approach and by integrating sociobiological perspectives. It aims at deciphering how individuals move, eat, care for and work and combines previous and new dataset on bioanthropology, bioge...

Programme scientifique - Avignon Université; 1. Une problématique : le vivant en tension |Le vivant nous interroge dans sa tension permanente avec les dynamiques culturelles, politiques, artistiques, économiques et technologiques qui le traversent. Constamment soumis à de nombreuses pressions externes qui peuvent le déformer, le vivant en tension peut être admis comme un mouvement perpétuel de résistance et de négociation, où sa perception et son rôle dans la société sont constamment redéfinis. En conséquence, la culture politique des corps enveloppe les chairs organiques, lunettes performatives et altérantes d’une dimension vivante à dévoiler. Dans un monde de plus en plus numérisé et rationalisé, l’intersection entre la création et le corps vivant donne les moyens d’étudier les enjeux de la communication humaine. Cette rencontre peut nous aider à mieux comprendre les enjeux de domination, transformation et réduction de l’organisme. Pour ce faire, nous avons investi le champ de la recherche-création, en invitant les chercheurs de diverses disciplines et objets de recherche à apporter leur contribution à cette problématique complexe. Des expressions classiques de l’écriture et du langage aux outils de réalité virtuelle, nous explorons aussi bien les nouvelles formes de pédagogie, que les expériences et représentations du vivant. Ce projet collectif permet ainsi de développer une meilleure compréhension de cette tension multifactorielle.2. Une méthodologie : la recherche-création |Notre pratique de la recherche-création vise à mêler recherche et création au sein d’un même processus méthodologique à l’université. La recherche-création fabrique des liens entre la création scientifique et artistique de manière à proposer une exploration organique de notre problématique du “vivant en tension”. Ce cadre de travail articule état de l’art, analyse de corpus et enquête anthropologique à une pratique créative qui s’incarne dans un organisme (chercheur, doctorant, étudiant). Dès lors, une réflexion sur les écritures sensibles peut s’opérer, tout comme d’autres projets scientifiques engagés dans une approche affective et singulière. La recherche-création permet également de répondre aux enjeux contemporains à travers plusieurs leviers pédagogiques : - la création comme initiation à la recherche ;- la création comme transmission des connaissances ;- la recherche-création comme méthode qualitative ;- la recherche-création comme médiation des univers artistiques et culturels.Notre approche a déjà fait l’objet de publications (Renucci et Galli, 2022 ; Galli et Rivière, 2022 ; Galli, 2021) qui ont permis de nourrir la réflexion sur l’offre de formation en SIC du Master Culture et Communication d’Avignon Université : nouveau module disponible en “recherche-création numérique” depuis 2022. 3. Un socle épistémologique : les séminaires de création | Ce programme de recherche ( https://recherche-creation.fr/ ) s’appuie sur un socle de plusieurs séminaires interdisciplinaires (ouverts aux chercheurs, doctorants, étudiants, enseignants, etc.) qui visent à explorer le vivant en tension avec la création : - Le séminaire “Créations inter-masters” (N. Macé-Barbier, D. Galli, G. Rivière) aborde la question des écritures du vivant auprès d’étudiants de différentes promotions de master (théâtre et communication), et avec le soutien d’artistes et d’établissements culturels ;- Le séminaire “Corps vivants et création” (V. Bénéï, D. Galli, G. Rivière, P. Hert) met en place une série d’expérimentations scientifiques dans le cadre de la recherche-création à partir du mouvement et de l’exploration des corps vivants. Ce séminaire implique Avignon Université, l’EHESS et Aix-Marseille Université ;- Le séminaire “Création-Médiation et réalité virtuelle” (C. Garson, D. Galli, G. Rivière) vise à interroger les applications vivantes de la réalité virtuelle dans les champs de l’enseignement et de la création numérique ;- Le séminaire “Création scientifique et communication” (F. Renucci, D. Galli, G. Rivière) interroge les frontières épistémologiques des processus de création à l’université. Entre rencontres, conversations, émotions, fabrications ou échanges documentaires, les traces des relations humaines au cœur de la recherche-création permettent d’approcher une nouvelle histoire anthropologique et communicationnelle du savoir. Enfin, ce programme de recherche vise à proposer un encadrement scientifique et artistique à des étudiants de master ou doctorat qui souhaitent réaliser un manuscrit préparé sous le Label Recherche-Création de l’École Doctorale 537 « Culture et Patrimoine » d’Avignon Université.

International audience; This exploratory study proposes an original intra-life history investigation through sequential analysis of the isotopic composition of carbon, nitrogen, and sulfur (CNS) on both primary and secondary dentin of a tooth (M1). We focus on an elderly woman from Longobard Italy (6th to 8th c. CE), who showed an unprecedented case of cranial surgery, presented in a companion paper by Micarelli and colleagues. Sequential stable CNS isotope composition of first molar dentin collagen allows us to infer diet, mobility, health, and physiological stress between approximately 3 months after birth to 9.5 years old (primary dentin) and between early adulthood until death (secondary dentin). Isotopic results on primary dentin highlight the following: (i) a long weaning period (ending at approximately 4 years), followed by (ii) a specific diet, including the contribution of C4 crops in early childhood (approximately 5.5 years), possibly concomitant with mobility. While secondary dentin shows a generally homogeneous diet during adulthood, the longitudinal analysis provided information on specific stresses that likely occurred in periods of difficult health conditions. This work emphasizes the importance of measuring complete dentin sequences (including secondary when present) for isotopic analysis to broaden intra-life histories in ancient populations.

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Ancient northeast Arabia underwent a phase of settlement expansion, as the region became a commercial crossroads between South Arabia, the Indian Ocean, Mesopotamia and the Mediterranean world. This "renaissance" is reflected in the emergence of vast tumulus necropolises that are part of a burial tradition dating back to the 3rd millennium BC. The most important of the sites founded at this time is Thaj. It is located in the present-day Eastern Province of Saudi Arabia. Explored within the framework of a Franco-Saudi archaeological mission, it consists of a fortified city of 40 ha, suburbs and a vast necropolis composed of more than 1000 funerary monuments, the archaeological and anthropological study of which is the focus of this doctoral thesis. The objectives of this research are to characterise the funerary practices and to fully understand the long occupation of this necropolis, from the first half of the 1st millennium BC to the 4th century AD.In order to answer these questions, the study was carried out according to three main research areas by applying a multidisciplinary approach. Firstly, through surveys, the spatial analysis of the necropolis allowed us to understand its organisation, its limits and its chronology. Secondly, archaeological excavations were carried out to establish a typology of the burial mounds and to reconstruct the rituals associated with the burial of the deceased. Finally, the osteological study of the skeletal remains was conducted to characterise the modalities of recruitment of the deceased within the necropolis.Based on the funerary data presented in this thesis, it is now possible to identify the first milestones in the evolution of funerary practices at Thaj and, more broadly, in northeast Arabia., Durant l’Antiquité, l’Arabie du Nord-Est connaît une phase d’expansion du peuplement, alors que la région devient un carrefour commercial entre l’Arabie du Sud, l’océan Indien, la Mésopotamie et le monde méditerranéen. Cette « renaissance » se traduit notamment par l’apparition de vastes nécropoles tumulaires qui s’inscrivent dans une tradition funéraire remontant au IIIe millénaire av. n. è. Le plus important des sites fondés à cette époque est Thaj. Il se trouve dans l’actuelle Province Orientale de l’Arabie Saoudite. Exploré dans le cadre d’une mission archéologique franco-saoudienne, il se compose d’une ville fortifiée de 40 ha, de faubourgs et d’une vaste nécropole composée de plus de 1000 monuments funéraires dont l’étude archéologique et anthropologique constitue le centre de cette thèse doctorale. Les objectifs de cette recherche sont de caractériser les pratiques funéraires et appréhender en totalité la longue occupation de cette nécropole, de la première moitié du Ier millénaire av. n. è. au IVe siècle de n. è.Pour répondre à ces problématiques, l’étude a été menée selon trois grands thèmes de recherche en appliquant une approche pluridisciplinaire. Premièrement, grâce aux prospections, l’analyse spatiale de la nécropole a permis de comprendre son organisation, ses limites et sa chronologie. Deuxièmement, des fouilles archéologiques ont été menées pour établir une typologie des tumulus et restituer les gestes associés à l’inhumation des défunts. Enfin, l’étude ostéologique des restes squelettiques a été conduite pour caractériser les modalités de recrutement des défunts au sein de la nécropole.À partir des données funéraires présentées dans cette thèse, il est aujourd’hui possible de dresser les premiers jalons de l’évolution des pratiques funéraires de Thaj et plus largement en Arabie du Nord- Est.

National audience; Les typo-chronologies sont utilisées fréquemment pour la datation des sépultures par les ­professionnels qui officient en archéologie funéraire. Elles concernent les éléments constitutifs des tombes : positions des défunts, contenants, rituels, aménagements et architectures funéraires de surface ou interne à la fosse. D’autres approches et méthodes y sont associées (radiocarbone, assemblages de mobilier, stratigraphie…).Les référentiels typo-chronologiques utilisés aujourd’hui sont parfois anciens et les avancées récentes de la recherche ont considérablement renouvelé la documentation en archéologie funéraire. Il a donc semblé nécessaire au Groupe d’Anthropologie et d’Archéologie Funéraire (Gaaf) de proposer de nouveaux référentiels en ce domaine à travers cet outil à destination des archéologues et des anthropologues. Il concerne les structures funéraires à inhumation, pour des périodes s’inscrivant dans un large horizon chronologique depuis la Préhistoire jusqu’à la période contemporaine. Cet ouvrage présente 46 articles, classés par grandes aires géographiques et issus des communications données lors de la 11e Rencontre du Gaaf, tenue à Tours du 3 au 5 juin 2019.

International audience; The sites of the PACA region have often been dated using the only two typochronological references available, established by Sylvain Gagnière (1965) and Michel Colardelle (1983 and 1996). Their use persisted despite their obsolescence highlighted during their application. The multiplication of new data, combined with methodological progress in the archaeology of death, enables the use of a new rich regional frame of reference from Antiquity to the modern period. This work aims to provide a relevant and appropriate tool for this region. The new typochronology should facilitate the understanding of funerary practices according to their specific context and enable both temporalities and regional developments to be addressed.; Les sites funéraires en PACA ont souvent été datés sur la base des deux seuls référentiels typochronologiques disponibles, établis par Sylvain Gagnière (1965) et Michel Colardelle (1983 et 1996). Leur utilisation a persisté malgré leur obsolescence souvent relevée. La multiplication de nouvelles données, associée aux progrès méthodologiques de l’archéologie de la Mort, a conduit à la mise en oeuvre d’un nouveau référentiel régional, de l’Antiquité à la période contemporaine. Ce travail a l’ambition de fournir un outil pertinent et adapté pour la région PACA. La nouvelle typo-chronologie proposée devrait faciliter l’appréhension des faits et des ensembles funéraires en fonction de leur contexte propre et permettre d’aborder aussi bien les temporalités et que les évolutions régionales.

International audience; In recent years, numerous authors have studied the joint attendance of shrines by worshippers of different religions or denominations in the Mediterranean region. In the last twenty years, this topic has become a kind of research sub-field that has produced a multitude of publications, collective research programmes, conferences and seminars. It has also been covered in media other than classical academic production, through exhibitions, films and websites. This first issue of "Religiographies. Representations, Texts and Lives" intends to contribute to this rapidly growing field.; Ces dernières années, de nombreux auteurs ont étudié la fréquentation conjointe, dans le monde méditerranéen, des sanctuaires par des fidèles de différentes religions ou confessions. Au cours des vingt dernières années, ce thème est devenu une sorte de sous-champ de recherche qui a donné lieu à une multitude de publications, de programmes collectifs de recherche , de conférences et de séminaires. Il a également été traité par des médias autres que la production académique classique, à travers des expositions, des films et des sites web. Ce premier numéro de "Religiographies. Representations, Texts and Lives" entend contribuer à ce domaine de recherche en pleine expansion.

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International audience; Childhood is an ontogenetic stage unique to the modern human life history pattern. It enables the still dependent infants to achieve an extended rapid brain growth, slow somatic maturation, while benefitting from provisioning, transitional feeding, and protection from other group members. This tipping point in the evolution of human ontogeny likely emerged from early Homo. The GAR IVE hemi-mandible (1.8 Ma, Melka Kunture, Ethiopia) represents one of the rarely preserved early Homo infants (~ 3 years at death), recovered in a richly documented Oldowan archaeological context. Yet, based on the sole external inspection of its teeth, GAR IVE was diagnosed with a rare genetic diseaseamelogenesis imperfecta (AI)-altering enamel. Since it may have impacted the child's survival, this diagnosis deserves deeper examination. Here, we reassess and refute this diagnosis and all associated interpretations, using an unprecedented multidisciplinary approach combining an in-depth analysis of GAR IVE (synchrotron imaging) and associated fauna. Some of the traits previously considered as diagnostic of AI can be better explained by normal growth or taphonomy, which calls for caution when diagnosing pathologies on fossils. We compare GAR IVE's dental development to other fossil hominins, and discuss the implications for the emergence of childhood in early Homo. The pattern and timing of modern human life history is unique in involving an extended period of growth 1. This provides more time not only for somatic development and protracted brain growth 2, 3 , but also to learn survival skills 4. The offspring is thus dependent upon more parental care over a longer time, and sexual maturity is postponed 5. In modern humans, life history stages successively involve infancy, childhood, a juvenile phase, adolescence, and finally adulthood. Different kinds of feeding strategies and dental development stages characterize these phases. Following these criteria, infancy can be subdivided into two stages. First, the "nursing phase" takes place from birth to 6 months, when the infant exclusively relies on breastfeeding and starts erupting its

Objective We report a probable case of multiple skeletal dysplasia observed in a Late Iron Age young adult male. Materials The individual studied belongs to a Late Iron Age necropolis from Switzerland. The skeletal elements are well preserved Methods Macroscopic and radiographic assessment. Results The individual shows evidence of both craniofacial and mandibular deformation. Developmental defects are also visible with effects on the general shape and articular surfaces of both humeri, as well as the left femur and tibia. Conclusion We propose that the lesions observed are manifestations of skeletal dysplasia, such as pseudo-achondroplasia or multiple epiphyseal dysplasia. Significance This is the first recorded case of multiple skeletal dysplasia in an Iron Age necropolis in Switzerland, questioning the integration of physically compromised individuals in La Tene society. Limitations Examination of other skeletal dysplasias from archaeological contexts provides support for this diagnosis. Suggestions for future research The integration of individuals with disabilities in La Tene societies is still poorly understood and further research is needed to better characterize these communities.

Figurative drawing is a skill that takes time to learn, and it evolves during different childhood phases that begin with scribbling and end with representational drawing. Between these phases, it is difficult to assess when and how children demonstrate intentions and representativeness in their drawings. The marks produced are increasingly goal-oriented and efficient as the child's skills progress from scribbles to figurative drawings. Pre-figurative activities provide an opportunity to focus on drawing processes. We applied fourteen metrics to two different datasets (N = 65 and N = 344) to better understand the intentional and representational processes behind drawing, and combined these metrics using principal component analysis (PCA) in different biologically significant dimensions. Three dimensions were identified: efficiency based on spatial metrics, diversity with color metrics, and temporal sequentiality. The metrics at play in each dimension are similar for both datasets, and PCA explains 77% of the variance in both datasets. Gender had no effect, but age influenced all three dimensions differently. These analyses for instance differentiate scribbles by children from those drawn by adults. The three dimensions highlighted by this study provide a better understanding of the emergence of intentions and representativeness in drawings. We discussed the perspectives of such findings in comparative psychology and evolutionary anthropology.

Gigantopithecus blacki was a giant hominid that inhabited densely forested environments of Southeast Asia during the Pleistocene epoch1. Its evolutionary relationships to other great ape species, and the divergence of these species during the Middle and Late Miocene epoch (16-5.3 million years ago), remain unclear2,3. Hypotheses regarding the relationships between Gigantopithecus and extinct and extant hominids are wide ranging but difficult to substantiate because of its highly derived dentognathic morphology, the absence of cranial and post-cranial remains1,3-6, and the lack of independent molecular validation. We retrieved dental enamel proteome sequences from a 1.9-million-year-old G. blacki molar found in Chuifeng Cave, China7,8. The thermal age of these protein sequences is approximately five times greater than that of any previously published mammalian proteome or genome. We demonstrate that Gigantopithecus is a sister clade to orangutans (genus Pongo) with a common ancestor about 12-10 million years ago, implying that the divergence of Gigantopithecus from Pongo forms part of the Miocene radiation of great apes. In addition, we hypothesize that the expression of alpha-2-HS-glycoprotein, which has not been previously observed in enamel proteomes, had a role in the biomineralization of the thick enamel crowns that characterize the large molars in Gigantopithecus9,10. The survival of an Early Pleistocene dental enamel proteome in the subtropics further expands the scope of palaeoproteomic analysis into geographical areas and time periods previously considered incompatible with the preservation of substantial amounts of genetic information. E.C. and F.W. are supported by VILLUM FONDEN (17649) and by the European Commission through a Marie Skłodowska-Curie (MSCA) Individual Fellowship (795569). T.M.-B. is supported by BFU2017-86471-P (MINECO/FEDER, UE), NIHM grant U01 MH106874, Howard Hughes International Early Career grant, Obra Social ‘La Caixa’ and Secretaria d’Universitats i Recerca and CERCA Programme del Departament d’Economia i Coneixement de la Generalitat de Catalunya (GRC 2017 SGR 880). E.C., J.C., J.V.O., D.S. and P.G. are supported by the Marie Skłodowska-Curie European Training Network (ETN) TEMPERA, a project funded by the European Union’s EU Framework Program for Research and Innovation Horizon 2020 under grant agreement no. 722606. M.J.C. and M.M. are supported by the Danish National Research Foundation award PROTEIOS (DNRF128). Work at the Novo Nordisk Foundation Center for Protein Research is funded in part by a donation from the Novo Nordisk Foundation (NNF14CC0001). Research at Chuifeng Cave is made possible by support from the National Natural Science Foundation of China (41572023) and by a grant from the Bagui Scholar of Guangxi. M.K. was supported by a Deutsche Forschungsgemeinschaft (DFG) fellowship (KU 3467/1-1) and the Postdoctoral Junior Leader Fellowship Programme from ‘la Caixa’ Banking Foundation (LCF/BQ/PR19/11700002). M.E.A. is supported by the Independent Research Fund Denmark (7027-00147B). We thank E. Willerslev for critical reading of the manuscript, scientific support and guidance.

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