Search

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.

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.).

The evolution and development of human mortuary behaviors is of enormous cultural significance. Here we report a richly-decorated young infant burial (AVH-1) from Arma Veirana (Liguria, northwestern Italy) that is directly dated to 10,211–9910 cal BP (95.4% probability), placing it within the early Holocene and therefore attributable to the early Mesolithic, a cultural period from which well-documented burials are exceedingly rare. Virtual dental histology, proteomics, and aDNA indicate that the infant was a 40–50 days old female. Associated artifacts indicate significant material and emotional investment in the child’s interment. The detailed biological profile of AVH-1 establishes the child as the earliest European near-neonate documented to be female. The Arma Veirana burial thus provides insight into sex/gender-based social status, funerary treatment, and the attribution of personhood to the youngest individuals among prehistoric hunter-gatherer groups and adds substantially to the scant data on mortuary practices from an important period in prehistory shortly following the end of the last Ice Age.

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.

The eastern Alpine region of Italy represents a well-known area with regards to the study of Late Pleistocene and Early Holocene forager adaptations in the whole peninsula. Caves, shelters and open-air sites, which are numerous in this region, have yielded traces of human re-peopling from mountaineous environments since the Late Glacial and until the Early Holocene. However, the archaeological record of the region lacks evidence about long lasting forager occupations. Exceptionally for the eastern Alpine sector, Riparo Biarzo (Natisone Valley, Julian Prealps) has yielded important evidence of continuous forager frequentation from Late Pleistocene to Early Holocene. The authors discuss the results of the analyses of animal remains and shell ornaments recovered in the Late Epigravettian and Mesolithic units of the site and their significance for understanding continuity and changes in the subsistence and symbolic strategies during these periods in the region. Archaeological data indicate an abundance of wild boar remains during both the Late Pleistocene and the Early Holocene at Biarzo, which makes hunting strategies at this site unique with regards to other contemporary contexts of the eastern Alpine region. Also, modalities of selection, production and use of ornaments in the same periods reveal the existence of established exchange networks, mobility strategies and connectivity between the eastern Alpine and the North Adriatic regions during the Late Epigravettian and throughout the Mesolithic.

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.

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.

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.

Cave bear ( Ursus spelaeus ), brown bear ( Ursus arctos ), and Neanderthals were potential competitors for environmental resources (shelters and food) in Europe. In order to reinforce this view and contribute to the ongoing debate on late Neanderthal behavior, we present evidence from zooarchaeological and taphonomic analyses of bear bone remains discovered at Rio Secco Cave and Fumane Cave in northeast Italy, an extended geographic area north of the Adriatic Sea. The remains from both caves come from layers dated to 49-42 ky cal. BP, and suggest close interactions between humans and bears, with data not only limited to the association of Mousterian lithic artifacts with numerous bear remains, but also the detection of clearly preserved traces of human modification such as cut and percussion marks, which enable a reconstruction of the main steps of fur recovery and the butchering process. Examples of Neanderthal bear exploitation are extremely sporadic in Europe, and Grotta Rio Secco and Grotta Fumane can be considered rare cases of remain accumulations generated by the human predation of bears of varied age classes during or near the end of hibernation. All of this evidence suggests that bears had a strategic role in the nomadic economy of Neanderthal hunting groups.

In attempt to enlighten the debate on the Middle to Upper Palaeolithic transition, this work contributes new data from a yet unpublished site, Riparo Broion in the north-east of Italy. Studies confirm the presence of the Uluzzian technocomplex embedded in an archaeological sequence which includes Mousterian, Gravettian and Epigravettian. These layers have yielded finds of bone and lithic technology, shell beads, engraved portable art and the use of red mineral pigments, which make this archive a unique case for evaluating the implications of cultural dynamics in Northern Italy 44.0–42.4ky BP. The diversity of the faunal assemblage recovered in layers 1f and 1g is representative of the different environments surrounding the site, with ungulates being targeted for hunting and consumption. The lithic assemblage records a high fragmentation rate due to the extensive use of the bipolar knapping technique, responsible for a large variety of splintered pieces and associated chips which also affected the backed pieces, lunates and end-scrapers. The features of the bone tools, as well as those of the marine and freshwater beads, echo the technique used in the Uluzzian in the south of Italy, confirming the geographic spread of this technocomplex in the North Adriatic region, as previously envisaged at Grotta Fumane. However, comparisons between these two sites stimulate a discussion around the possible driving forces responsible for the differences observed in technological, typological and structural lithic assemblages. A definitive scenario for the Middle Palaeolithic–Upper Palaeolithic transition in this region of Southern Europe is ongoing and Riparo Broion is adding key contributions.

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.

Hammerstones and anvils are among the oldest tools used by hominins to perform a variety of tasks including knapping activities. The bipolar technique on anvil is well documented in Prehistory since the Lower Palaeolithic and is usually considered to be an expedient technique in comparison to other knapping systems. This technique plays a pivotal role in the Uluzzian techno-complex lithic production where it is largely used. In the present study, we analyse the anvils and hammerstones recovered in the Mousterian and Uluzzian layers of the site of Castelcivita (Campania region-southern Italy) by a multi-disciplinary approach. Our aim is to investigate the function and functioning of anvils and hammerstones by evaluating the presence and the role of bipolar knapping in these two assemblages. To do this, we integrated techno-functional analysis (sensu Boeda) and use-wear study, by defining each techno-functional unit (transformative and prehensile unities) of anvils and hammerstones and identifying the specific use-wear left by the bipolar technique by means of a dedicated experimental programme. The obtained results allowed us to observe different technical behaviours, concerning both the production and the use of hammerstones and anvils, between Mousterian and Uluzzian. Differences were encountered in the selection of raw material (limestone in the Uluzzian, sandstone in the Mousterian) and in the technical way of approaching the tool as well as in the function: the Mousterian anvil was used ‘as is’ for crushing materials, unlike the Uluzzian anvils which were exclusively employed for bipolar knapping, after adapting their original volume. Hammerstones were mainly used as pestles or retouchers in the Mousterian and for direct percussion in knapping activities during the Uluzzian.