A ZooMS-informed archaeozoological and taphonomic analysis comparing Neanderthal and Homo sapiens subsistence behaviours in Northwest Italy

Ce projet contribue aux discussions en cours sur la transition du Paléolithique moyen au Paléolithique supérieur en Europe occidentale marquée par un tournant dans l’évolution de notre espèce, l’Homo sapiens. Alors que les Néandertaliens, nos plus proches cousins évolutionnaires disparaissent du registre fossile, les humains modernes qui ont migré hors d’Afrique, se dispersent rapidement à travers l’Eurasie. Les deux populations étaient exposées aux mêmes changements climatiques dramatiques caractéristiques de la transition, et pourtant, les Néandertaliens sont rapidement remplacés par les humains modernes. Par conséquent, ce phénomène suggère que les populations humaines modernes auraient pu être mieux adaptées face aux changements environnementaux. Puisque le régime alimentaire est un bon moniteur de l’adaptation, cette recherche compare les stratégies de subsistance des deux espèces humaines ayant tour à tour occupé le site de Riparo Bombrini (Balzi Rossi, Ligurie, Italie). Une analyse archéozoologique et taphonomique a été effectuée sur les collections fauniques du Moustérien tardif et du Proto-Aurignacien afin d’obtenir la première comparaison détaillée du régime alimentaire et des comportements de chasse des Néandertaliens et des humains modernes sur l’un des seuls sites du nord-ouest de l’Italie entièrement documenté avec des méthodes archéologiques modernes. Étant donné que la nature très fragmentée des ossements animaux sur le site a été un obstacle aux analyses fauniques dans le passé, les méthodes d’analyse archéozoologique ont été complétées par le « collagen fingerprinting » (c.-à-d. zooarchéologie par spectrométrie de masse, ou ZooMS) afin d’assurer l’identification d’un maximum de spécimens pour atteindre une précision accrue de l’identification taxonomique. La préservation différentielle du collagène dans les restes squelettiques a également justifié le développement d’une méthode novatrice de dépistage du collagène utilisant la spectroscopie FTIR-ATR pour la présélection d’échantillons ZooMS. Les résultats montrent que, tandis que Néandertal et Homo sapiens ont continuellement chassé les taxons ongulés disponibles à proximité de Riparo Bombrini, les niveaux de Moustérien tardif indiquent un rétrécissement du tableau de chasse associé à un mode de subsistance hyperlocal. En revanche, les spectres fauniques se sont considérablement élargis dans le plus ancien Proto-Aurignacien, lorsque Riparo Bombrini était occupé comme camp de base logistique à long terme associé à un vaste territoire de subsistance. Les résultats fournissent également les premières données détaillées sur la subsistance des populations humaines durant la transition dans la région de l’arc liguro-provençal, établissant ainsi de nouvelles hypothèses à tester dans de futurs travaux concernant la nature changeante de leurs écologies.This project contributes to the ongoing debates over the Middle-Upper Paleolithic transition in Western Europe, which marks a turning point in the evolution of our species, Homo sapiens. While Neanderthals, our closest evolutionary relatives, went extinct at that time, modern humans who had migrated out of Africa dispersed very rapidly across Eurasia. While both populations were exposed to the same dramatic climatic shifts at the time, it is only the Neanderthals that quickly disappeared from the archeological record, suggesting that modern human populations may have been better adapted to react to environmental changes than Neanderthals. Since diet is a good monitor of adaptation, this research compares the subsistence strategies of both human groups as they occupied, in quick succession, the site of Riparo Bombrini (Balzi Rossi, Liguria, Italy). An archeozoological and taphonomic analysis was conducted on Late Mousterian and Proto-Aurignacian faunal collections to produce the first direct comparison between Neanderthal and modern human diets and hunting strategies at one of the only sites in Northwest Italy entirely excavated using modern documentation methods. Because the highly fragmented nature of the animal bones at the site has hindered faunal analysis in the past, these approaches were complemented by collagen fingerprinting (i.e., Zooarcheology by Mass Spectrometry, or ZooMS) to identify as many specimens as possible as to species, thus yielding unprecedented accuracy in taxonomic identification. The challenging collagen preservation state also required developing a screening method using FTIR-ATR spectroscopy prior to ZooMS. The results show that, while Neanderthals and modern humans continuously hunted prime-aged ungulate taxa available in a close range of Riparo Bombrini, the Late Mousterian levels indicate a narrower diet associated with a hyper-local subsistence range. In contrast, the faunal spectra broadened noticeably in the earliest Proto-Aurignacian, when Riparo Bombrini was occupied as a long-term logistical base camp within an extensive land-use strategy. The results also provide the first high-resolution view of human subsistence during the transition in the Liguro-Provençal arc region and set up test hypotheses about the changing nature of hominin behavioural ecology that can be further tested in future work

Grotte du Bison : deux chasseurs pour un gibier : analyse archéozoologique de la couche I-J sur le site moustérien de la Grotte du Bison, Arcy-sur-Cure (Yonne, France)

Le site moustérien de la Grotte du Bison est situé au cœur des grottes préhistoriques longeant la rivière de la Cure à Arcy-sur-Cure (Yonne, France). La couche I-J de ce gisement représente une occupation néandertalienne datant de la fin du stade isotopique 4 (MIS 4) et du début du stade 3. Face à l’instabilité du climat durant cette période, les groupes de chasseurs-cueilleurs néandertaliens doivent faire des choix en matière d’acquisition des ressources alimentaires. Le mode de subsistance de ces groupes implique des choix stratégiques de comportements de chasse, de transport et de traitement des carcasses, ainsi que de mobilité de groupe. Quelles sont les stratégies d’exploitation de la faune employées par les Néandertaliens à Arcy-sur-Cure il y a un peu plus de 50 000 ans? Ce mémoire présente une analyse archéozoologique de l’assemblage faunique mis au jour durant la mission de fouilles 2014. Les résultats indiquent que les populations néandertaliennes ont occupé le site de façon saisonnière en alternance avec d’autres animaux carnivores comme l’ours des cavernes et la hyène des cavernes. Les hyènes et les Néandertaliens sont deux potentiels agents accumulateurs d’ossements dans la grotte. Un regard taphonomique sur l’assemblage faunique de la couche I-J suggère que les groupes néandertaliens ont chassé le renne et le cheval, alors que les meutes de hyènes ont accumulé des ossements de bovinés et de chevaux. Les groupes néandertaliens de la Grotte du Bison ont rapporté les carcasses entières de leurs proies sur le site. Ils en ont exploité la viande, la moelle, les peaux, ont fabriqué des outils en os et ont utilisé les plumes des rapaces, vraisemblablement à des fins symboliques.The “Grotte du Bison” is a Mousterian site located along the river Cure at Arcy-sur-Cure (Yonne, France). Level I-J is a Neanderthal occupation of the cave that occurred during the end of marine isotope stage 4 (MIS 4) and the beginning of stage 3. This period is characterised by strong climate instability, which conditioned the Neanderthals’ choices regarding their subsistence. Subsistence patterns are defined by various behaviours such as the type of hunting strategy adopted, carcass transport and butchery practices and residential mobility. What subsistence strategies were adopted by Neanderthal groups who occupied the Grotte du Bison over 50 000 years ago? This thesis presents a zooarchaeological analysis of the faunal assemblage excavated during the 2014 excavations. The results indicate that Neanderthal populations seasonally occupied the site alternating with carnivores such as bears (hibernating in the cave during the winter) and hyenas. Both Neanderthals and hyenas served as bone accumulators in the cave. Preliminary taphonomic analysis of the faunal assemblage from level I-J suggest that the Neanderthals hunted horses and reindeers while hyenas hunted and/or scavenged bovines and horses. Neanderthals from the Grotte du Bison transported whole carcasses to the site where they butchered them, exploiting their bone marrow and grease, processed the skins, made bone tools, and used raptor feathers (possibly as items of personal adornment)

An infant burial from Arma Veirana in northwestern Italy provides insights into funerary practices and female personhood in early Mesolithic Europe

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

Un outil en os à usages multiples dans un contexte moustérien

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Archaeozoological, Taphonomic and ZooMS Insights into The Protoaurignacian Faunal Record from Riparo Bombrini

Human adaptation to climatic variations is being discussed at different scales and from diverse perspectives and specializations in Paleolithic archaeology. We suggest examining human mobility on the local scale through the faunal record to better understand human-environmental interactions during the early dispersal of anatomically modern humans along the Mediterranean coast. Riparo Bombrini is located in the renowned Balzi Rossi complex in Northwest Italy. The site offers an excellent opportunity to compare two distinct Protoaurignacian levels yielding well-documented and well-dated deposits. Previous studies of spatial, lithic, and raw material data from these two Protoaurignacian levels have revealed distinct mobility signatures as well as undeniable evidence for the resilience of the Protoaurignacian technocomplex during episodes of climatic instability including the HE4 event, circa 40ka cal BP. The highly fragmented nature of the animal bones at the site warrants the application of the ZooMS (Zooarchaeology by Mass Spectrometry) collagen fingerprinting technique. For this research we carried out taphonomic and archaeozoological analyses with integrated systematic ZooMS using a mass sampling strategy. The results suggest stability in hunting strategies over time in spite of the apparent shift in mobility strategies from level A2 to level A1 at Riparo Bombrini

Portable FTIR for on-site screening of archaeological bone intended for ZooMS collagen fingerprint analysis

Faunal remains play an important role in helping reconstruct Palaeolithic hunter-gatherer subsistence and mobility strategies. However, differential bone preservation is an issue in southern European prehistoric sites, which often makes morphological identification impossible. Zooarchaeology by Mass Spectrometry (ZooMS) is a new, low-cost method that can improve NISP statistical significance in a replicable way by using diagnostic peptides of the dominant collagen protein as a fingerprint of animal (including hominin) species. It is also a powerful tool to assess collagen preservation for radiocarbon dating. This paper presents the proof of concept of a method for evaluating collagen preservation in a quick and minimally destructive way in the field prior to ZooMS analysis by using a portable Fourier Transform Infrared Spectrometer (FTIR) equipped with an attenuated total reflectance accessory (ATR). The method was tested on faunal assemblages from two north-western Italian sites: Riparo Bombrini and Arma Veirana. Both are important sites for understanding the Middle and Upper Palaeolithic of Liguria but are located in two distinct environmental contexts (coast vs. mountainous hinterland) that impacted collagen preservation. The relative abundance of collagen in powdered bone samples was evaluated by calculating the Amide I to phosphate ratio (CO/P) from spectra collected with two portable and three laboratory-based FTIR instruments. The bones were then analysed by ZooMS and the results were compared to evaluate the effectiveness of using FTIR as a screening technique. Results indicate that FTIR instruments are excellent tools to predict collagen preservation in fragmented archaeological bones, but a screening method should be first calibrated on the analysed faunal assemblage by conducting a test of the CO/P screening procedure with various measurement systems on a single FTIR instrument

Risk and resilience in the late glacial: A case study from the western Mediterranean.

The period spanning the Last Glacial Maximum through early Holocene encompasses dramatic and rapid environmental changes that offered both increased risk and new opportunities to human populations of the Mediterranean zone. The regional effects of global climate change varied spatially with latitude, topography, and distance from a shifting coastline; and human adaptations to these changes played out at these regional scales. To better understand the spatial and temporal dynamics of climate change and human social-ecological-technological systems (or SETS) during the transition from full glacial to interglacial, we carried out a meta-analysis of archaeological and paleoenvironmental datasets across the western Mediterranean region. We compiled information on prehistoric technology, land-use, and hunting strategies from 291 archaeological assemblages, recovered from 122 sites extending from southern Spain, through Mediterranean France, to northern and peninsular Italy, as well as 2,386 radiocarbon dates from across this region. We combine these data on human ecological dynamics with paleoenvironmental information derived from global climate models, proxy data, and estimates of coastlines modeled from sea level estimates and digital terrain. The LGM represents an ecologically predictable period for over much of the western Mediterranean, while the remainder of the Pleistocene was increasingly unpredictable, making it a period of increased ecological risk for hunter-gatherers. In response to increasing spatial and temporal uncertainty, hunter-gatherers reorganized different constituents of their SETS, allowing regional populations to adapt to these conditions up to a point. Beyond this threshold, rapid environmental change resulted in significant demographic change in Mediterranean hunter-gatherer populations

Machine learning ATR-FTIR spectroscopy data for the screening of collagen for ZooMS analysis and mtDNA in archaeological bone

Faunal remains from archaeological sites allow for the identification of animal species that enables the better understanding of the relationships between humans and animals, not only from their morphological information, but also from the ancient biomolecules (lipids, proteins, and DNA) preserved in these remains for thousands and even millions of years. However, due to the costs and efforts required for ancient biomolecular analysis, there has been considerable research into development of accurate and efficient screening approaches for archaeological remains. FTIR spectroscopy is one such approach that has been considered for screening of proteins, but its widespread use has been hindered by the fact that its predictive accuracy can vary widely depending on the extent of sample preservation and the instrument used. Further, screening methods for ancient DNA (aDNA) analysis are scarce. Here we present a new approach to vastly improve upon FTIR-based screening methods prior to ZooMS (Zooarchaeology by Mass Spectrometry) and aDNA analysis through the use of random forest-based machine learning. To do so, we use ATR-FTIR to examine three sets of archaeological bone assemblages and analyse them by ZooMS (for taxonomic identification). Two of these are from Palaeolithic contexts, dominated by terrestrial fauna and include specimens with a variety of preservational conditions. The third set consists of Holocene faunal remains, with variable levels of preservation and is dominated by cetaceans. Using the Holocene faunal remains, we were able to more consistently evaluate ATR-FTIR-based screening for mtDNA as well as ZooMS success. We report on the potential of machine learning in ATR-FTIR-based screening for ancient mtDNA analysis, and our machine learning models conclusively improve the accuracy prior to usage of ATR-FTIR-based screening for ZooMS by 20\u201340%. The results also suggest this approach potentially allows for a universal screening system, applicable across multiple sites and largely independent of the spectrometers used

Machine Learning ATR-FTIR Spectroscopy Data for the Screening of Collagen for ZooMS Analysis and mtDNA in Archaeological Bone

Supplementary Tables for the article 'Machine Learning ATR-FTIR Spectroscopy Data for the Screening of Collagen for ZooMS Analysis and mtDNA in Archaeological Bone' for presentation with the University of Manchester Thesis 'Analysis of Proteins from Archaeological Artefacts, with a Focus on Ancient Human Diet

An infant burial from Arma Veirana in northwestern Italy provides insights into funerary practices and female personhood in early Mesolithic Europe

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