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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.

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 (

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.

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.

Come e dove hanno vissuto i Neandertal? Cosa ha provocato la loro scomparsa? Queste domande intrattengono la comunità scientifica da molti anni, in un dibattito che ha visto il susseguirsi delle ipotesi più svariate: dai cambiamenti climatici alla convivenza e/o competizione con i Sapiens per l’occupazione del territorio e lo sfruttamento delle risorse, fino a ipotizzare devastanti effetti tumorali, legati alle oscillazioni del campo magnetico terrestre, ovviamente a scapito dei nativi. Negli ultimi decenni grande attenzione viene posta sulla possibile esistenza di qualche forma di contatto tra gli ultimi Neandertal e i primi Uomini Anatomicamente Moderni (Homo sapiens), giunti nel sud dell'Europa circa 45mila anni fa. Il bacino del Mediterraneo e, con esso, le zone settentrionali come l'alto Adriatico e la fascia delle Prealpi si pongono al centro di questo dibattito. Le culture e i comportamenti che hanno modulato le strategie adattative dell’umanità del passato, si sono tradotti in varie attività connesse alla pura sopravvivenza o alle pratiche cultuali. Ciò è testimoniato dalla diversità di elementi archeologici, il cui potenziale di informazioni culturali permette di collegare tra loro i siti archeologici e le conoscenze acquisite con la loro esplorazione. Nella regione dell'Adriatico settentrionale, in particolare nelle Prealpi, ampie documentazioni archeologiche consentono di ricostruire il modo di vita degli ultimi Neandertal e di comprendere come questi, nel corso di centinaia di miglia di anni siano stati in grado di adattarsi a numerosi mutamenti climatici e ambientali. L’elevata disponibilità di materia prima litica e la varietà fisiografica ed ecologica della frangia al cardine tra l’alta pianura e le Prealpi spiegano l’interesse alla frequentazione di questi territori, percorsi in spostamenti stagionali. In relazione alla loro posizione topografica, alcune grotte principali delle Prealpi Venete (Grotta di Fumane, Riparo Tagliente, Grotta di San Bernardino) fungevano da siti di riferimento, a frequentazione complessa, dove la produzione di attrezzi in pietra scheggiata era finalizzata alla caccia e al processamento delle prede. A questi contesti si associavano siti complementari a frequentazione breve ed effimera, talora prossimi a fonti primarie di approvvigionamento minerale e faunistico, e utilizzati strategicamente negli ampi percorsi logistici. In questi casi, il frazionamento delle sequenze di produzione/trasformazione evidenziato dalle industrie litiche rivela la capacità di pianificare le attività economiche e la variabilità nella circolazione dei manufatti. Uno di questi casi si trova nelle Prealpi Carniche, un distretto sino a pochi anni fa poco noto all'archeologia preistorica fino a due decenni fa. Si tratta della Grotta del Rio Secco, un'ampia cavità sull'Altopiano di Pradis, il cui archivio stratigrafico delinea un potenziale scientifico come indicatore archeologico di occupazione del territorio, di comportamento alimentare, di sfruttamento delle risorse minerali in un settore chiave tra la pianura e la regione alpina. Unitamente al transetto pedemontano circostante, l’Altopiano di Pradis cade all’interno di una provincia interposta tra tanto diverse quanto complementari entità fisiografiche ed ecologiche: l’area veneta a occidente, ricca di siti all’aperto e di grotte e ripari; le Alpi Giulie, dove si apre la Grotta Divje Babe I; il carso Triestino con vari siti in grotta; la Croazia nordoccidentale e la costa dalmata, che ospitano giacimenti noti ma anche di recente scoperta. Grotta del Rio Secco è stata per noi e per tutti i nostri collaboratori una grande occasione di crescita: sul piano scientifico, consentendoci di esplorare aspetti del comportamento di una specie umana del passato poco noti alla comunità scientifica internazionale; sul piano sociale, costruendo una profonda rete di contatti e amicizie, che ha reso questa esperienza solida e ineguagliabile.

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.

The avian fossil assemblages from the late Pleistocene deposits of the Rio Secco Cave (north-eastern Italy) is presented herein. We studied the layers that date back to the end of MIS3 and the beginning of MIS2, which also contain evidence of Gravettian frequentation dated to 33.5–30 ka cal BP. The systematic analysis revealed the presence of 18 species and other supraspecific taxa that supported palaeoenvironmental reconstructions. Taxa indicate that, at the onset of LGM, site surroundings were characterised by conifer or mixed forests, open grasslands, slow-flowing water bodies and mountain meadows with rocky outcrops, as indicated by Lagopus muta. Today, this environment is found above the tree line (beyond 1500–2000 m) and cannot be detected near the site, located at 580 m asl. Noteworthy, is also the finding of the second Italian late Pleistocene fossil record of Picus canus.

Objectives The aim of the study is the assessment of Nadale 1, a Neanderthal deciduous tooth recently discovered in Northeastern Italy in the De Nadale cave (Middle Palaeolithic). Together with the clear archaeological context of the site, this study brings new insight on Neanderthal behavior and dental morphological variability. Materials and methods We used microCT data to provide a morphological description and morphometric analysis (diameter measurements and dental tissue volumes) of the Nadale 1 human tooth. Microwear analysis, taphonomical investigation and caries identification were performed using a stereomicroscope and Scanning Electron Microscope. Results In terms of morphology (i.e., incipient tuberculum molare, marked mesial marginal ridge and well-developed mid-trigonid crest connecting the protoconid and the metaconid, deep anterior fovea) and size, Nadale 1 presents features frequently observed in Neanderthal lower first deciduous molars. Microscope investigations reveal the presence of a small pit which could be correlated to an incipient caries. Conclusion Nadale 1 expands the Italian Middle Palaeolithic fossil record and provides further information on Neanderthal dm1s in terms of dimensional and morphological variability. Furthermore, the presence of an incipient caries brings further data on Neanderthal diet.

The search for diagnostic criteria useful in hunting lesions identification is a new branch of investigation. Though recently there has been an increase in studies focused on this issue, no experimental works exist that analyze marks left by backed, morphologically standardized lithic projectiles like those used by the hunter-gatherers that peopled a large part of Europe during the Late Glacial. As such, this paper aims to provide comparison data for identifying archaeological Late Epigravettian projectile impact marks. At the same time, the potential of 3D scanning microscopy to distinguish hunting injuries from other taphonomic marks is assessed. The morphometric analyses, based on the descriptive criteria developed from other recent studies, highlight the presence of peculiar features of experimentally produced drag and puncture marks. These data are interpreted as a result of the specific design of Late Epigravettian lithic projectiles. The outcomes of 3D digital analysis confirm the crucial role of this methodological approach in taphonomic study, offering new clues in PIMs (Projectile Impact Marks) archaeological identification and distinction from cut marks, carnivore tooth marks and corrosion cavities.

From the intricate ensemble of evidence related to the Middle–Upper Palaeolithic transition and the presumed first spread of anatomically modern humans in Europe, the Uluzzian has attracted major attention in the past few years. Although the Uluzzian has been supposedly viewed as a product of modern humans settling in Mediterranean Europe, the techno-cultural complex has been the subject of few investigations aiming to clarify its chronology, bone industry, and settlement dynamics. Further, little is known of its technological structure. This article presents the results of an extensive study of the lithic and bone technologies from assemblages recovered at Fumane Cave in the north of Italy. Results confirm that the Uluzzian is a flake-dominated industry that brings together a set of technological innovations. The Levallois is the most used method in the initial phase, which is replaced by more varied flaking procedures and an increase in bladelets and flake-blades. Sidescrapers and points also represent a Mousterian feature in the initial phase, while splintered pieces, backed knives and other Upper Palaeolithic tools increase in the later phase. Our results suggest that the Uluzzian is rooted in the Mousterian lithic technological context and cannot be viewed as a proxy for anatomically modern humans, the carriers of the abrupt cultural changes related with the Aurignacian.

We present the results of a multi-disciplinary investigation on a deciduous human tooth (Pradis 1), recently recovered from the Epigravettian layers of the Grotte di Pradis archaeological site (Northeastern Italian Prealps). Pradis 1 is an exfoliated deciduous molar (Rdm2), lost during life by an 11—12-year-old child. A direct radiocarbon date provided an age of 13,088—12,897 cal BP (95% probability, IntCal20). Amelogenin peptides extracted from tooth enamel and analysed through LC¿MS/MS indicate that Pradis 1 likely belonged to a male. Time-resolved 87Sr/86Sr analyses by laser ablation mass spectrometry (LA-MC-ICPMS), combined with dental histology, were able to resolve his movements during the first year of life (i.e. the enamel mineralization interval). Specifically, the Sr isotope ratio of the tooth enamel differs from the local baseline value, suggesting that the child likely spent his first year of life far from Grotte di Pradis. Sr isotopes are also suggestive of a cyclical/seasonal mobility pattern exploited by the Epigravettian human group. The exploitation of Grotte di Pradis on a seasonal, i.e. summer, basis is also indicated by the faunal spectra. Indeed, the nearly 100% occurrence of marmot remains in the entire archaeozoological collection indicates the use of Pradis as a specialized marmot hunting or butchering site. This work represents the first direct assessment of sub-annual movements observed in an Epigravettian hunter-gatherer group from Northern Italy., The authors are grateful to Municipal Administration of Clauzetto, Pradis Cave Museum and the Superintendence for Archeology, Fine Arts and Landscape of Friuli Venezia Giulia for granting permission to study the tooth. 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.; www.erc-success.eu and Grant Agreement no. 803147–RESOLUTION awarded to S.T.; https://site.unibo.it/resolution-erc/en) and by the MIUR FARE programme 2018 (FARE Ricerca in Italia: Framework per l'attrazione e il rafforzamento delle eccellenze—SAPIENS project to S.B.). A.N. is supported by a Marie Skłodowska-Curie Actions Individual Fellowship (grant number H2020-MSCA-IF-2018-842812-WEAN IT). E.Ba. is supported by EQUIPEX ASTER-CEREGE and ANR MARCARA and thanks Y. Fagault and T. Tuna for technical support. The Frankfurt Isotope & Element Research Center (FIERCE) is financially supported by the Wilhelm and Else Heraeus Foundation and by the Deutsche Forschungsgemeinschaft (DFG; INST 161/921-1 Forschungsgroßgeräte and INST 161/923-1 Forschungsgroßgeräte), which is gratefully acknowledged. This is FIERCE contribution No. 91. We acknowledge Elettra Sincrotrone Trieste for providing access to its laboratory facilities and the TomoLab staff for technical support. The authors deeply thank prof. Anna Cipriani for the use of geochemical facilities at UNIMORE (https://www.geochem.unimore.it/). Dr. Filippo Genovese is thanked for the help during LC-MS analyses. The ‘Fondazione Cassa di Risparmio di Modena’ is acknowledged for funding the UHPLC-ESI-Q Exactive system at the Centro Interdipartimentale Grandi Strumenti (CIGS), UNIMORE. Two anonymous reviewers are acknowledged for their constructive comments on our manuscript.