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This study is part of an ongoing project aiming to unravel the different phases of growth during talar development. Overall, our preliminary results suggest that age-related morphological variations of the talus may be used to determine the general age of juvenile skeletal remains, which could be valuable to many archaeological and forensic researchers. Future studies will explore larger samples, including individuals younger than 12 months, and a more in depth analysis (i.e., single surfaces) to better evaluate differences between groups.

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During the restoration of the Conversion of San Paul’s church, in Roccapelago (Italy), a hidden crypt was brought to light. Therein, a large amount of human skeletal remains was retrieved, including a considerable amount of disarticulated non-adult specimens, belonging to the inhabitants that lived there from the last decades of the 16th century to the end of the 18th century. Simultaneously, parish registers of birth and death were found and digitized. This is the first study focused on the juvenile post medieval community of Roccapelago, which aims to provide new data about infant mortality and paleopathology during the 16th and 18th centuries, through the comparison of anthropological data to information available in parish records. Standard anthropological protocols were used to assess the Minimum Number of Individuals, age-at-death and pathologies. Results showed that at least 161 non-adults were buried into the crypt. The mortality range was high among perinates, especially between the 36th and the 40th weeks (26%), and during the first postnatal year, particularly in the first six months (11%). Then, mortality rates fell within the 5th years (2.4%). Parish records confirmed the high mortality rates at birth and among the first postnatal year, linked to the risks associated to the birth and the peril of the weaning period. The pathological analysis highlighted the presence of metabolic diseases, such as scurvy. This study provides a unique opportunity to compare anthropological protocols for age estimation to the information registered in the parish records when dealing with commingled juvenile remains.

Acknowledgements: The authors thank G. Marciani and O. Jöris for comments on archaeology; C. Jeong, M. Spyrou and K. Prüfer for comments on genetics; M. O’Reilly for graphical support for Fig. 5 and Extended Data Fig. 9; the entire IT and laboratory teams at the Department of Archaeogenetics of MPI-SHH for technical assistance; M. Meyer and S. Nagel for support with single-stranded library preparation; K. Post, P. van Es, J. Glimmerveen, M. Medendorp, M. Sier, S. Dikstra, M. Dikstra, R. van Eerden, D. Duineveld and A. Hoekman for providing access to human specimens from the North Sea (The Netherlands); M. D. Garralda and A. Estalrrich for providing access to human specimens from La Riera (Spain); J. Górski and M. Zając for providing access to human specimens from Maszycka cave; C. Di Patti for providing access to human specimens from San Teodoro 2 (Italy); P. Blaževičius for providing access to the Donkalnis human remains and the new radiocarbon dates; the Italian Ministry of Culture and Soprintendenza Archeologia Belle Arti e Paesaggio for the Provinces of Verona, Rovigo, and Vicenza for granting access to the human remains of Tagliente 2; F. Fontana, who carries out investigations of the Riparo Tagliente site (Italy); the Friuli Venezia Giulia Superintendency for providing access to the human tooth Pradis 1; and the Soprintendenza Archeologia Belle Arti e Paesaggio for the Provinces of Barletta-Andria-Trani and Foggia for providing access to the Paglicci human remains. This project has received funding by the European Research Council under the European Union’s Horizon 2020 research and innovation programme under grant agreements no. 803147-RESOLUTION (to S.T.), no. 771234-PALEoRIDER (to W.H.), no. 864358 (to K.M.), no. 724703 and no. 101019659 (to K.H.). K.H. is also supported by the Deutsche Forschungsgemeinschaft (DFG FOR 2237). E.A. has received funding from the Van de Kamp fonds. PACEA co-authors of this research benefited from the scientific framework of the University of Bordeaux’s IdEx Investments for the Future programme/GPR Human Past. A.G.-O. is supported by a Ramón y Cajal fellowship (RYC-2017-22558). L. Sineo, M.L. and D.C. have received funding from the Italian Ministry of University and Research (MUR) PRIN 2017 grants 20177PJ9XF and 20174BTC4R_002. H. Rougier received support from the College of Social and Behavioral Sciences of CSUN and the CSUN Competition for RSCA Awards. C.L.S. and T. Saupe received support from the European Union through the European Regional Development Fund (project no. 2014-2020.4.01.16-0030) and C.L.S. received support from the Estonian Research Council grant PUT (PRG243). S. Shnaider received support from the Russian Science Foundation (no. 19-78-10053)., Modern humans have populated Europe for more than 45,000 years1,2. Our knowledge of the genetic relatedness and structure of ancient hunter-gatherers is however limited, owing to the scarceness and poor molecular preservation of human remains from that period3. Here we analyse 356 ancient hunter-gatherer genomes, including new genomic data for 116 individuals from 14 countries in western and central Eurasia, spanning between 35,000 and 5,000 years ago. We identify a genetic ancestry profile in individuals associated with Upper Palaeolithic Gravettian assemblages from western Europe that is distinct from contemporaneous groups related to this archaeological culture in central and southern Europe4, but resembles that of preceding individuals associated with the Aurignacian culture. This ancestry profile survived during the Last Glacial Maximum (25,000 to 19,000 years ago) in human populations from southwestern Europe associated with the Solutrean culture, and with the following Magdalenian culture that re-expanded northeastward after the Last Glacial Maximum. Conversely, we reveal a genetic turnover in southern Europe suggesting a local replacement of human groups around the time of the Last Glacial Maximum, accompanied by a north-to-south dispersal of populations associated with the Epigravettian culture. From at least 14,000 years ago, an ancestry related to this culture spread from the south across the rest of Europe, largely replacing the Magdalenian-associated gene pool. After a period of limited admixture that spanned the beginning of the Mesolithic, we find genetic interactions between western and eastern European hunter-gatherers, who were also characterized by marked differences in phenotypically relevant variants.

This project aims to test whether geometric morphometric (GM) and trabecular analyses may be useful tools in identifying talar characteristics related to hypochondroplasia. We quantified the external and internal talar morphology of an hypochondroplasic dwarf (T17) from Modena (northern Italy) dated to the 6th century AD. External talar morphology of T17 was compared with a broad sample of modern human tali (n = 159) using GM methods. Additionally, a subsample of these tali (n = 41) was used to investigate whole talar trabecular changes in T17. Our results show that GM and trabecular analyses identify a combination of traits linked to the dwarfing disorder of hypochondroplasia. These traits include decreased scaled talar dimensions compared to normal-sized individuals, presence of an accessory antero-lateral talar facet, high bone volume fraction and high anisotropy values throughout the entire talus. In our case study, hypochondroplasia does not appear to substantially modify external talar morphology probably due to the fast growth of the talus. We suggest that small talar dimensions are associated with hypochondroplasia. An antero-lateral talar facet may result from the talus and calcaneus coalition (i.e., talocalcaneal abnormal bridging) possibly related to an everted foot posture that was limited by overgrowth of the fibula. We suggest that high talar trabecular density and strut orientation provide insights into pathological development of the trabecular plates in T17. Finally, our study suggests that high talar trabecular density and strut orientation, and small talar dimensions, may be added as possible concomitant talar hallmarks for hypochondroplasia.

The reconstruction of the original morphology of bones and teeth after sampling for physicochemical (e.g., radiocarbon and uranium series dating, stable isotope analysis, paleohistology, trace element analysis) and biomolecular analyses (e.g., ancient DNA, paleoproteomics) is appropriate in many contexts and compulsory when dealing with fossil human remains. The reconstruction protocols available to date are mostly based on manual re-integration of removed portions and can lead to an imprecise recovery of the original morphology. In this work, to restore the original external morphology of sampled teeth we used computed microtomography (microCT), reverse engineering (RE), computer-aided design (CAD) and rapid prototyping (RP) techniques to fabricate customized missing parts. The protocol was tested by performing the reconstruction of two Upper Palaeolithic human teeth from the archaeological excavations of Roccia San Sebastiano (Mondragone, Caserta, southern Italy) and Riparo I of Grotte Verdi di Pradis (Clauzetto, Pordenone, north-eastern Italy) (RSS2 and Pradis 1, respectively), which were sampled for physicochemical and biomolecular analyses. It involved a composite procedure consisting in: a) the microCT scanning of the original specimens; b) sampling; c) the microCT scanning of the specimens after sampling; d) the reconstruction of the digital 3D surfaces of the specimens before and after sampling; e) the creation of digital models of the missing/sampled portions by subtracting the 3D images of the preserved portions (after the sampling) from the images of the intact specimens (before the sampling) by using reverse engineering techniques; f) the prototyping of the missing/sampled portions to be integrated; g) the painting and application of the prototypes through the use of compatible and reversible adhesives. By following the proposed protocol, in addition to the fabrication of a physical element which is faithful to the original, it was possible to obtain a remarkable correspondence between the contact surfaces of the two portions (the original and the reconstructed one) without having to resort to any manipulation/adaptation of either element., This project was funded by the European Research Council (ERC) under the European Union’s Horizon 2020 Research and Innovation Programme (Grant Agreement no. 724046–SUCCESS awarded to S.B.)

Objectives The study of the development of human bipedalism can provide a unique perspective on the evolution of morphology and behavior across species. To generate new knowledge of these mechanisms, we analyze changes in both internal and external morphology of the growing human talus in a sample of modern human juveniles using an innovative approach. Materials and Methods The sample consists of high-resolution microCT scans of 70 modern juvenile tali, aged between 8 postnatal weeks and 10 years old, from a broad chronological range from Middle/Late Neolithic, that is, between 4800 and 4500 BCE, to the 20th century. We applied geometric morphometric and whole-bone trabecular analysis (bone volume fraction, degree of anisotropy, trabecular number, thickness, and spacing) to all specimens to identify changes in the external and internal morphology during growth. Morphometric maps were also generated. Results During the first year of life, the talus has an immature and globular shape, with a dense, compact, and rather isotropic trabecular architecture, with numerous trabeculae packed closely together. This pattern changes while children acquire a more mature gait, and the talus tends to have a lower bone volume fraction, a higher anisotropy, and a more mature shape. Discussion The changes in talar internal and external morphologies reflect the different loading patterns experienced during growth, gradually shifting from an “unspecialized” morphology to a more complex one, following the development of bipedal gait. Our research shows that talar plasticity, even though genetically driven, may show mechanical influences and contribute to tracking the main locomotor milestones.

Objective: The development of bipedalism is a very complex activity that contributes to shaping the anatomy of the foot. The talus, which starts ossifying in utero, may account for the developing stages from the late gestational phase onwards. Here, we explore the early development of the talus in both its internal and external morphology to broaden the knowledge of the anatomical changes that occur during early development.Materials and Methods: The sample consists of high-resolution microCT scans of 28 modern juvenile tali (from 36 prenatal weeks to 2 years), from a broad chronological range from the Late Roman period to the 20th century. We applied geometric morphometric and whole-bone trabecular analysis to investigate the early talar morphological changes.Results: In the youngest group (

Neanderthal extinction has been a matter of debate for many years. New discoveries, better chronologies and genomic evidence have done much to clarify some of the issues. This evidence suggests that Neanderthals became extinct around 40,000–37,000 years before present (BP), after a period of coexistence with Homo sapiens of several millennia, involving biological and cultural interactions between the two groups. However, the bulk of this evidence relates to Western Eurasia, and recent work in Central Asia and Siberia has shown that there is considerable local variation. Southwestern Asia, despite having a number of significant Neanderthal remains, has not played a major part in the debate over extinction. Here we report a Neanderthal deciduous canine from the site of Bawa Yawan in the West-Central Zagros Mountains of Iran. The tooth is associated with Zagros Mousterian lithics, and its context is preliminary dated to between ~43,600 and ~41,500 years ago.

Before the end of the Last Glacial Maximum (LGM, ∼16.5 ka ago) set in motion major shifts in human culture and population structure, a consistent change in lithic technology, material culture, settlement pattern, and adaptive strategies is recorded in Southern Europe at ∼18–17 ka ago. In this time frame, the landscape of Northeastern Italy changed considerably, and the retreat of glaciers allowed hunter-gatherers to gradually recolonize the Alps. Change within this renewed cultural frame (i.e., during the Late Epigravettian phase) is currently associated with migrations favored by warmer climate linked to the Bølling-Allerød onset (14.7 ka ago), which replaced earlier genetic lineages with ancestry found in an individual who lived ∼14 ka ago at Riparo Villabruna, Italy, and shared among different contexts (Villabruna Cluster). Nevertheless, these dynamics and their chronology are still far from being disentangled due to fragmentary evidence for long-distance interactions across Europe. Here, we generate new genomic data from a human mandible uncovered at Riparo Tagliente (Veneto, Italy), which we directly dated to 16,980–16,510 cal BP (2σ). This individual, affected by focal osseous dysplasia, is genetically affine to the Villabruna Cluster. Our results therefore backdate by at least 3 ka the diffusion in Southern Europe of a genetic component linked to Balkan/Anatolian refugia, previously believed to have spread during the later Bølling/Allerød event. In light of the new genetic evidence, this population replacement chronologically coincides with the very emergence of major cultural transitions in Southern and Western Europe., The research was supported by the European Union through the European Research Council under the European Union’s Horizon 2020 Research and Innovation Programme (grant agreement no. 724046 – Success awarded to S.B., http://www.erc-success.eu; grant agreement no. 803147 Resolution awarded to S.T., https://site.unibo.it/resolution-erc/en) as well as through the European Regional Development Fund (project no. 2014–2020.4.01.16–0030 to C.L.S. and T.S.) and projects no. 2014-2020.4.01.16-0024 and MOBTT53 (L.P.), by the Estonian Research Council personal research grant (PRG243; C.L.S.), and by UniPd PRID 2019 (L.P.).

Strontium isotopes are applied to a wide range of scientific fields and to different types of sample materials, providing valuable information foremost about provenance and age, but also on diagenetic processes and mixing relationships between different Sr reservoirs. The development of in-situ analytical techniques, such as laser ablation ICP-MS, has improved our understanding of Sr isotope variability in several field of application, because of the possibility to discriminate small-scale changes and their spatial distribution. However, large outputs of Sr isotope data are produced by laser ablation MC-ICP-MS systems, which necessitate of multiple offline steps to correct and assess the data. This requires the availability of simple and user-friendly tools, easily manageable also by non-specialists. With this in mind, we developed SrDR, an Excel-based interactive data reduction spreadsheet (‘SrDR’, Sr-Data-Reduction) for the processing of Sr isotopes measured by LA-MC-ICP-MS. The SrDR spreadsheet is easily customizable (a) to meet user-specific analytical protocols, (b) for different instruments (i.e. Nu Plasma vs. Neptune), and (c) for diverse target materials (e.g. Rare Earth Elements enriched or depleted samples). We include also several examples relevant to low and high temperature geochemistry fields - a fossil tooth, a modern seashell, a speleothem sample and plagioclase crystals - to show how different sample materials are corrected for different interfering masses.

The early onset of weaning in modern humans has been linked to the high nutritional demand of brain development that is intimately connected with infant physiology and growth rate. In Neanderthals, ontogenetic patterns in early life are still debated, with some studies suggesting an accelerated development and others indicating only subtle differences vs. modern humans. Here we report the onset of weaning and rates of enamel growth using an unprecedented sample set of three late (���70 to 50 ka) Neanderthals and one Upper Paleolithic modern human from northeastern Italy via spatially resolved chemical/isotopic analyses and histomorphometry of deciduous teeth. Our results reveal that the modern human nursing strategy, with onset of weaning at 5 to 6 mo, was present among these Neanderthals. This evidence, combined with dental development akin to modern humans, highlights their similar metabolic constraints during early life and excludes late weaning as a factor contributing to Neanderthals��� demise

Roccia San Sebastiano is a tectonic-karstic cave located at the foot of the southern slope of Mt. Massico, in the territory of Mondragone (Caserta) in Campania (southern Italy). Systematic excavation has been carried out since 2001, leading to the partial exploration of an important Pleistocene deposit, extraordinarily rich in lithic and faunal remains. The aim of this paper is to (1) present the stratigraphic sequence of Roccia San Sebastiano, and (2) technologically describe the lithic materials of squares F14 t18, t19, t20; E16 t16, t17, t18 recently recognised as Uluzzian. The stratigraphic sequence is more than 3 m thick and dates from the Middle to the Upper Palaeolithic. It contains different techno-complexes: Gravettian, Aurignacian, Uluzzian and Mousterian. In the Uluzzian lithic assemblage mostly local pebbles of chert were used in order to produce small-sized objects. The concept of debitage mainly deals with unidirectional debitage with absent or fairly accurate management of the convexities and angles; the striking platforms are usually natural or made by one stroke. It is attested the use of both direct freehand percussion and bipolar technique on anvil in the same reduction sequence. Amongst the retouched tools the presence of two lunates is of note. This study of the Roccia San Sebastiano Uluzzian lithic complexes is significant for understanding the dynamics of the transition from Middle to Upper Palaeolithic in the Tyrrhenian margin of southern Italy.

Many aspects of diet and behaviour can be gleaned from dental wear, including environmental conditions, food-fracture properties, food processing techniques, and cultural habits. Specifically, the angulation (flat vs. steep) of molar wear has been used as an indicator of food toughness and has also been implicated in the use of grinding stones and pottery in food processing. In this work, we focus on the sequential phases of the power stroke of mastication in a functional way, measuring the inclination of molar wear facets through the occlusal fingerprint analysis method. Specifically, we have calculated the angulation of wear facets in upper and lower molars of Palaeolithic humans, extant hunter-gatherers, and proto-farmers to discern differences between groups with different diets. Contrary to previous analyses, our study shows that the molars of Late Pleistocene specimens are characterized by significantly steeper angles than those of modern hunter-gatherers. The flat molar wear found in the latter group could be related to the excessive mixture of exogenous materials accidentally introduced into their foods, as indicated by ethnographic evidence. On the contrary, the steep wear angles characterizing the Palaeolithic group are probably associated with the consumption of a less abrasive diet, which could be ultimately due to food preparation techniques that incorporated less dust and grit into their diets.

The late Middle Palaeolithic (MP) settlement patterns in the Levant included the repeated use of caves and open landscape sites. The fossil record shows that two types of hominins occupied the region during this period—Neandertals and Homo sapiens. Until recently, diagnostic fossil remains were found only at cave sites. Because the two populations in this region left similar material cultural remains, it was impossible to attribute any open-air site to either species. In this study, we present newly discovered fossil remains from intact archaeological layers of the open-air site ‘Ein Qashish, in northern Israel. The hominin remains represent three individuals: EQH1, a nondiagnostic skull fragment; EQH2, an upper right third molar (RM3); and EQH3, lower limb bones of a young Neandertal male. EQH2 and EQH3 constitute the first diagnostic anatomical remains of Neandertals at an open-air site in the Levant. The optically stimulated luminescence ages suggest that Neandertals repeatedly visited ‘Ein Qashish between 70 and 60 ka. The discovery of Neandertals at open-air sites during the late MP reinforces the view that Neandertals were a resilient population in the Levant shortly before Upper Palaeolithic Homo sapiens populated the region.

Defining the processes involved in the technical/cultural shifts from the Late Middle to the Early Upper Palaeolithic in Europe (~50-39 thousand years BP) is one of the most important tasks facing prehistoric studies. Apart from the technological diversity generally recognised as belonging to the latter part of the Middle Palaeolithic, some assemblages showing original technological traditions (i.e. Initial Upper Palaeolithic: Bohunician, Bachokirian; so called transitional industries: Châtelperronian, Szeletian, Lincombian-Ranisian-Jerzmanowician, Uluzzian; Early Upper Palaeolithic: Protoaurignacian, Early Aurignacian) first appear during this interval. Explaining such technological changes is a crucial step in order to understand if they were the result of the arrival of new populations, the result of parallel evolution, or of long-term processes of cultural and biological exchanges. In this debate Italy plays a pivotal role, due to its geographical position between eastern and western Mediterranean Europe as well as to it being the location of several sites showing Late Mousterian, Uluzzian and Protoaurignacian evidence distributed across the Peninsula. Our study aims to provide a synthesis of the available lithic evidence from this key area through a review of the evidence collected from a number of reference sites. The main technical features of the Late Mousterian, the Uluzzian and the Protoaurignacian traditions are examined from a diachronic and spatial perspective. Our overview allows the identification of major differences in the technological behaviour of these populations, making it possible to propose a number of specific working hypotheses on the basis of which further studies can be carried out. This study presents a detailed comparative study of the whole corpus of the lithic production strategies documented during this interval, and crucial element thus emerge: 1. In the Late Mousterian tools were manufactured with great attention being paid to the production phases and with great investment in inizializing and managing core convexities; 2. In contrast, Uluzzian lithic production proceeded with less careful management of the first phases of debitage, mainly obtaining tool morphologies by retouching. 3. In the Protoaurignacian the production is carefully organized and aimed at obtaining laminar blanks (mainly bladelets) usually marginally retouched. These data are of primary importance in order to assess the nature of the "transition" phenomenon in Italy, thus contributing to the larger debate about the disappearance of Neandertals and the arrival of early Modern Humans in Europe.

Human skeletal remains from archaeological contexts occasionally present signs of traumatic injuries from weapons, revealing, for example, the degree of interpersonal violence, the type of weapon and the sequence of events of a specific historical context. Traumatic lesions are generally analyzed using macroscopic and microscopic methods, which are not necessarily integrated in the same study. In this study, we employed a multi-analytical approach to determine if new, or more refined information could be gained compared to traditional analyses. Here, we describe and analyze interpersonal skeletal peri-mortem sharp-force trauma in remains recovered in 2009 during archaeological excavations of a cemetery in the 4th to 6th century AD the city of Modena (Italy). Evidence for sharp force trauma was observed in four of the 13 individuals recovered at site. The traumatic lesions were analyzed using an integrated multi-analytical approach that combines traditional macroscopic examination, light microscopy, and three-dimensional digital modeling. We aimed to determine the number, size, and the position of the lesions, and the direction (angles) of weapon penetration in the bone. In particular, we applied digital protocols for the orientation of the skeletal regions of interest involved in the trauma to provide useful results for future comparisons and investigations.

Objectives Exact symmetry and perfect balance between opposite jaw halves, as well as between antagonistic teeth, is not frequently observed in natural masticatory systems. Research results show that asymmetry in our body, skull, and jaws is often related to genetic, epigenetic, environmental and individual ontogenetic factors. Our study aims to provide evidence for a significant link between masticatory asymmetry and occlusal contact between antagonist teeth by testing the hypothesis that tooth inclination is one of the mechanisms driving distribution of wear in masticatory phases in addition to dietary and cultural habits. Materials and Methods The present work investigates the relationship between dental macrowear patterns and tooth inclinations on a sample of complete maxillary and mandibular 3D models of dental arches from 19 young and adult Yuendumu Aboriginal individuals. The analysis was carried out on first molars (M1) from all quadrants. Occlusal Fingerprint Analysis was used for the quantification of macrowear patterns, and 2D cross‐sectional geometric analysis was carried out to investigate asymmetry in dental arches. Results The asymmetry is highly variable on both arches, and it is associated with differences in the inclination of upper M1 crowns. Each molar has variable inclination (buccal/lingual) which influence tooth to tooth contact, producing greater or lesser variation in wear pattern. Interindividual variability of morphological variation of the occlusal relationship has to be considered in macrowear analysis. Discussion Our results suggest that overall asymmetry in the masticatory apparatus in modern humans affects occlusal contact areas between antagonist teeth influencing macrowear and chewing efficiency during ontogeny.

Here we report additional Upper Palaeolithic human remains from the Imereti region, western Georgia (SOM Fig. S1): two isolated teeth from Dzudzuana cave, Dzu 1 and Dzu 2 (both deciduous; Bar-Yosef et al., 2011), and one isolated tooth (SATP5-2, deciduous) and a hemi-mandible (SATP5) bearing permanent and deciduous teeth (SATP5-3 e SATP5-7) from Satsurblia cave (Pinhasi et al., 2014). In particular, the human remains from Dzudzuana cave, dated between 27,000 and 24,000 cal. BP, fill a huge gap in the Upper Palaeolithic Georgian fossil record and play an important role in the debate about modern human peopling of the Caucasus.