Human remains from Arma di Nasino (Liguria) provide novel insights into the paleoecology of early Holocene foragers in northwestern Italy.

We report the discovery and analysis of new Mesolithic human remains-dated to ca. 10,200-9000 cal. BP-from Arma di Nasino in Liguria, northwestern Italy, an area rich in Upper Paleolithic and Neolithic attestations, but for which little information on Early Holocene occupation was available. The multi-proxy isotopic profile of the two individuals reveals that-despite the proximity of the site to the Mediterranean seashore and the use of shellfish as decorative elements in burials-the ecology of these foragers was based on the exploitation of high-altitude resources, presumably in the nearby western Alps. This constitutes the first direct evidence in northwestern Italy of a significant ecological shift towards higher altitudes following deglaciation, especially when compared to isotopic data of the Late Pleistocene hunter-gatherers from the nearby site of Arene Candide Cave, who exploited terrestrial resources nearer to the coast and at lower altitudes. While the biochemistry of Nasino's skeletal assemblage revealed new details on Early Holocene lifeways in the area, the osteobiography of one individual offers glimpses into the life experience of a specific female forager, depicting a scenario of early skeletal trauma, developmental disturbances, long-term impairments, and resilience amongst the last European hunter-gatherers.

Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers.

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.

Multi-proxy analysis suggests Late Pleistocene affinities of human skeletal remains attributed to Balzi Rossi.

In two publications from 1967 and 1971, M. Masali described human skeletal remains presumed to have been found in the Balzi Rossi caves (Ventimiglia, Italy), based on a signed note dated to 1908. Since then, the remains - dubbed "Conio's Finds" and preserved at the University of Torino - had not been further studied. We performed a multidisciplinary investigation aimed at clarifying the geographical and chronological attribution of these specimens. Collagen extraction for AMS dating was unsuccessful, but we obtained two direct dates on the best- preserved crania via 231Pa/235U direct gamma-ray spectrometry (10,500±2,000 years BP and 12,500±2,500 years BP). We analyzed the metrics and morphology of the crania and femora by comparing them with samples belonging to the Upper Paleolithic, Mesolithic, and Neolithic periods, and evidenced that the "Conio's Finds" are morphologically more compatible with a Late Pleistocene rather than Holocene attribution. We analyzed the literature regarding the history of excavations at Balzi Rossi, and we propose that - if any credence should be given to the note accompanying the material - the remains may have been found in front of Grotta dei Fanciulli or Grotta del Caviglione, in the redeposited soil dug up during the installation of lime kilns carried out between the late 18th and the early 19th centuries. These hypotheses may be tested in the future by comparing the speleothem deposited on one of the crania and the remaining deposit at the site.

Tracking the transition to agriculture in Southern Europe through ancient DNA analysis of dental calculus.

Archaeological dental calculus, or mineralized plaque, is a key tool to track the evolution of oral microbiota across time in response to processes that impacted our culture and biology, such as the rise of farming during the Neolithic. However, the extent to which the human oral flora changed from prehistory until present has remained elusive due to the scarcity of data on the microbiomes of prehistoric humans. Here, we present our reconstruction of oral microbiomes via shotgun metagenomics of dental calculus in 44 ancient foragers and farmers from two regions playing a pivotal role in the spread of farming across Europe-the Balkans and the Italian Peninsula. We show that the introduction of farming in Southern Europe did not alter significantly the oral microbiomes of local forager groups, and it was in particular associated with a higher abundance of the species Olsenella sp. oral taxon 807. The human oral environment in prehistory was dominated by a microbial species, Anaerolineaceae bacterium oral taxon 439, that diversified geographically. A Near Eastern lineage of this bacterial commensal dispersed with Neolithic farmers and replaced the variant present in the local foragers. Our findings also illustrate that major taxonomic shifts in human oral microbiome composition occurred after the Neolithic and that the functional profile of modern humans evolved in recent times to develop peculiar mechanisms of antibiotic resistance that were previously absent.

Complex mortuary dynamics in the Upper Paleolithic of the decorated Grotte de Cussac, France.

The Mid-Upper Paleolithic (Gravettian) karstic Grotte de Cussac (France) contains two areas of human remains in the context of abundant (and spectacular) parietal engravings. The first area (loci 1 and 2) includes the skeleton of a young adult male in a bear nest, rearranged by postdecomposition inundation, and the variably fragmentary remains of at least two individuals distributed across two bear nests, sorted anatomically and with most of the elements constrained to one side of one nest. The second area (locus 3) retains remains of two adults and an adolescent, in upper hollows and variably distributed down the slope, largely segregated into upper versus lower body groups. The only decoration associated with the human remains is red pigment on some of the bones or underlying sediment. The human remains indicate variable nonnatural deposition and manipulation of human bodies, body portions, and skeletal elements of at least six individuals. Moreover, Cussac is unusual in the association of these remains with exceptional parietal art. The complex Cussac mortuary pattern joins growing evidence from other Gravettian sites of variable treatment of individuals after death, within and across sites, in terms of formal deposition of the body versus postmortem manipulation versus surface abandonment. It provides a window onto the social diversity and the complex interactions of the living and the dead among these successful Late Pleistocene foragers.

Insights on patterns of developmental disturbances from the analysis of linear enamel hypoplasia in a Neolithic sample from Liguria (northwestern Italy).

OBJECTIVE: To assess developmental disturbances through the analysis of linear enamel hypoplasia (LEH) frequency and to infer environmental stress and life history within Neolithic communities from Liguria (Italy). MATERIALS: 43 unworn/minimally worn permanent anterior teeth of 13 individuals recovered from nearby caves and dated to c. 4800-4400 cal. BCE. METHODS: LEH defects were identified with high-resolution macrophotos of dental replicas, age at LEH was calculated via perikymata counts. LEH defects matched between two or more teeth were considered as systemic disturbances. LEH frequency by age classes was analyzed via GLZ and Friedman ANOVA. RESULTS: Number of matched defects per individual range between 2-12. The mean LEH per individual was highest in the 2.5-2.99 age category, with a significant increase relative to earlier growth stages, followed by a decline. CONCLUSION: LEH may reflect life-history in the local ecology of Neolithic Liguria, where several individuals with osteoarticular tuberculosis have been recorded. Disease burden may have triggered developmental disturbances around the time of weaning. Age at first defect was negatively correlated with age at death and positively with the total number of defects, suggesting that early stress may have affected survivorship. SIGNIFICANCE: The study contributes to the reconstruction of ecological pressures among Neolithic people of Liguria, and informs on environmental challenges during the Neolithic adaptive expansion. LIMITATIONS: The visual examination of macrophotos is prone to observer error; mid-crown tends to display more visible LEH due to tooth architecture. SUGGESTIONS FOR FURTHER RESEARCH: Apply different quantitative methods to examine severity and duration of disturbances.

Morphological variability of Upper Paleolithic and Mesolithic skulls from Sicily.

Scenarios for the dispersal of Homo sapiens in Southern Europe and in the Mediterranean basin have been uncertain, given the scarceness of osteological samples and the simplicity of the proposed archaeologically-based settlement hypotheses. According to available data, the first anatomically modern humans entered Sicily during the Late Pleistocene, coming from the Italian peninsula. A presumably small Late Epigravettian population colonised coastal sites. Later, North-Western archaeological horizons gave hospitality to a significant Mesolithic expansion. In order to verify a hypothesis of continuity in the peopling of the island, we analyzed Sicilian skulls from the Late Epigravettian site of San Teodoro, Eastern Sicily (AMS 14C dated at 14,500 BP) and from the Mesolithic period (14C dated from 9,500 to 8,500 BP) coming from various sites (Uzzo, Molara, Grotta d'Oriente) located on the North Western coast of the island. The aims were to test the biological variability through time within the island as well as to evaluate the relationships of Sicilian Pleistocene hunter-gatherers with Old World populations. We also evaluated the Sicilian Mesolithic uniformity especially between the Uzzo and Grotta d'Oriente sites, given their vicinity and accessibility during the Early Holocene. We applied 3D geometric morphometric methods to assess shape variation as well as geographic and diachronic morphological patterns. All analyzed specimens, plus a comparative sample from the Old World dated from the Upper Paleolithic to recent periods, were transformed in digital images and standard craniofacial landmarks were extracted from the 3D models. Our results underline a high variability among the Mesolithic specimens, as well as a large craniometric distance from the presumed founder Paleolithic settler representatives (San Teodoro specimens) that have closer morphological affinities with other European Upper Paleolithic specimens.

New insights on Final Epigravettian funerary behavior at Arene Candide Cave (Western Liguria, Italy).

We gained new insights on Epigravettian funerary behavior at the Arene Candide cave through the osteological and spatial analysis of the burials and human bone accumulations found in the cave during past excavations. Archaeothanathological information on the human skeletal remains was recovered from diaries, field pictures and notes, and data from recent excavations was integrated. The secondary deposits have traditionally been interpreted as older burials that were disturbed to make space for new inhumations. Our results suggest that those disturbances were not casual: older burials were intentionally displaced to bury younger inhumations. Subsequently, some skeletal elements, especially crania, were arranged around the new burial; these were often placed within stone niches. Based on the composition of some clusters, which contain the bones of two individuals displaced together, it is possible that a double burial composed of two adults was originally present at the site. This would be a burial type that had not been recognized at Arene Candide until now. Strikingly, this potential double burial contained an individual showing pathological bowing of the limbs, a finding which is not infrequent in skeletons from Gravettian and Epigravettian multiple burials. In addition, the crania and other skeletal elements derived from this burial were intentionally placed around a new inhumation, whose skeleton possibly shows a milder form of the same disease (possibly hereditary rickets). This and other observations suggest that the five individuals belonging to the second phase of this "cemetery" (AMS dates spanning 12,030 -11,180 cal BP) might have been buried over a relatively brief time span. Our study demonstrated similar behaviors in the first phase of mortuary use of the cave (12,820-12,420 cal BP), indicating a remarkable persistence in Final Epigravettian funerary models despite their archaeologically apparent rarity and intermittent nature.

Smaller long bone cross-sectional size in people who died of tuberculosis: Insights on frailty factors from a 19th and early 20th century Finnish population.

There is little research on how individuals suffering from tuberculosis may differ from those not infected in terms of overall skeletal morphology. Tuberculosis was endemic in 19th and early 20th century Finland making documented skeletal collections of Finns ideal to study effects of the disease on bone. The present study compares long bone cross-sectional total area between individuals who died of tuberculosis and those with another recorded cause of death in a Finnish sample. Adult male individuals (N = 105) were selected for analysis. Complete humeri (N = 56), femora (N = 66) and tibiae (N = 64) were 3D scanned using a laser scanner and total cross-sectional areas calculated with AsciiSection software. Individuals who died of tuberculosis (N = 24, 15 humeri, 14 femora, 13 tibiae) had, when standardized for body size, significantly smaller total cross-sectional femoral and humeral, but not tibial, areas. The mechanisms behind the observed relationship may reflect a combination of biological 'frailty' in terms of susceptibility to infection, reduced childhood activity and/or vitamin D deficiency, which possibly influenced both subperiosteal development during adolescence and, later, susceptibility to contracting and dying of TB. Due to the relatively small sample future studies are needed to further investigate the relationship between TB and bone cross-sectional size.

Social reorganization and biological change: An examination of stature variation among Iron Age Samnites from Abruzzo, central Italy.

Stature is a sensitive indicator of overall environmental quality experienced during growth and development, and can provide insights on a population's 'well-being'. This study investigated changes in estimated adult stature in a large (N=568) sample of Samnite Iron Age (800-27 BCE) people from central Italy, during a period of increasing sociopolitical complexity. Stature was analyzed diachronically, between sexes, and across social strata inferred using the 'Status Index' based on funerary treatment. It was expected: 1) a decrease in stature from the Orientalizing-Archaic period (O-A) to the fifth century BC (V SEC) and the following Hellenistic period (ELL), due to population increase and urbanization; 2) social status to positively influence the attainment of the full stature potential; 3) sexual dimorphism to be higher in more stratified groups. Results revealed no significant diachronic changes in stature (females: O-A: 154.2cm,V SEC: 154.2cm, and ELL: 153.6cm; males: O-A: 165.0cm,V SEC: 165.2cm, and ELL: 165.0cm) or sexual dimorphism. High-status males were taller than low-status (p=0.021), possibly due to a better diet, but only in the Orientalizing-Archaic period. Nonsignificant changes in females suggest either differential access to resources in women, or a better buffering from environmental optima or crises. The results of this study highlight the complex interrelation between social factors and human growth, and stress the importance of understanding the specific mechanisms leading to variation in adult stature.

A 6500-year-old Middle Neolithic child from Pollera Cave (Liguria, Italy) with probable multifocal osteoarticular tuberculosis.

Clear skeletal evidence of prehistoric tuberculosis (TB) is rare, especially in children. We describe and differentially diagnose the pathological changes displayed by a five-year-old child, Pollera 21 (PO21) dated to the Middle Neolithic of Liguria (Italy), or 5740±30 BP (Beta-409341; 6635-6453cal BP, 2σ, OxCal 4.2). PO21 shows a number of osteoarticular lesions, mainly of a lytic nature with very little bone proliferation: the vertebral column, the shoulder and pelvic girdles, and the ribcage are involved. Given the nature and pattern of the lesions, we propose a diagnosis of multifocal (or multiple) bone TB. Attempts to detect TB aDNA through molecular analysis gave negative results, but this alone is not sufficient to prove that PO21 was not infected with TB. The lesions observed in PO21 share similarities with other published evidence, such as spinal and joint involvement, and disseminated cyst-like lesions. Conversely, PO21 does not show diffuse bone deposition, such as hypertrophic osteoarthropathy (HOA) or endocranial modifications such as serpens endocrania symmetrica (SES). PO21 adds to our knowledge of patterns of TB manifestation in archaeological skeletal remains, which is especially important considering the variability in types and patterns of osteoarticular lesions seen today in people with TB.

Insights on the paleoepidemiology of ancient tuberculosis from the structural analysis of postcranial remains from the Ligurian Neolithic (northwestern Italy).

The aim of this research is to gain insights on the progression timeline of osteoarticular tuberculosis (TB) in people from the Neolithic period by using skeletal traits that are independent of the bony lesions. The body proportions and postcranial mechanical strength of bones from two individuals from Liguria in northwestern Italy (Arene Candide 5, adolescent, and Arma dell'Aquila 1, adult), were compared with the rest of the Ligurian Neolithic skeletal series (45 individuals). If TB led to wasting of the skeleton and lack of normal function that endured for years, as often happens today, a clear signature of postcranial gracility and disruption of development should be apparent. Conversely, rapid progress of the disease would leave little systemic macroscopic change in the skeleton, except for the bony lesions directly caused by the TB pathogen, suggesting a different level of bacterial virulence in the past. The extreme biomechanical gracility observed in the lower limb of Arene Candide 5 suggests a period of compromised diaphyseal periosteal apposition during ontogeny due to metabolic disturbances likely linked to TB. Results suggest that, in Neolithic Liguria, TB in humans saw a slow, chronic progression, which is characteristic of diseases with long histories of host-pathogen co-evolution.

A bioarchaeological approach to the reconstruction of changes in military organization among Iron Age Samnites (Vestini) From Abruzzo, Central Italy.

The Samnites were an Iron Age population that shifted from warlike mountain dwellers to the largest sociopolitical unit of central Italy, able to dispute with Rome the domination over the peninsula. Archaeological and historical evidence suggests that this major shift in the scale of conflict may have involved a reorganization of the military system, which changed from an elite militia to a conscript or standing army from the Orientalizing-Archaic (800-500 BC) to Hellenistic times (400-27 BC). We propose a bioarchaeological framework jointly analyzing skeletal properties and funerary treatment in male Samnites to investigate on this shift in military organization. We anticipated that, when Samnites had an elite militia, the warring force was constituted by the wealthier segments of the society. Conversely, we expected the warring force of the standing/conscript army to be mainly drawn from the lower social strata. We considered high asymmetry in J, a measure of humeral torsional rigidity (calculated via cross-sectional geometry, CSG) as a proxy for pre- and peri-adolescent-onset weapon training. The social standing of the individual was approximated via funerary treatment analysis (Status Index). Results show that in the Orientalizing-Archaic period, humeral asymmetry and Status Index are positively correlated, and the high-status subsample shows significantly higher asymmetry than the low-status subsample. Among Hellenistic Samnites, no correlation between Status Index and humeral asymmetry is present, and the low-status subsample is the most lateralized. Results support the use of CSG in a strong theoretical framework to investigate past changes in military organization and their correlates in terms of sociopolitical development, alterations of power relationships, and warfare.

Mobility and subsistence economy: a diachronic comparison between two groups settled in the same geographical area (Liguria, Italy).

The purpose of this article is to investigate temporal shifts in skeletal robusticity to infer behavioral changes in two populations (Neolithic, NEOL and Medieval, MED) settled in the same geographic area but involved in different subsistence economies (pastoralism and coastal resources exploitation). This comparison allows us to test the hypothesis that occupational stress and mobility in the same environment produce predictable changes in the robusticity of both upper and lower limbs. Results show a lower degree of humeral robusticity and a similar degree of humeral asymmetry in the two sexes in the MED population. These results are consistent with the relatively less stressful subsistence economy documented in the MED population relative to that of the NEOL. Lower limb results suggest that femoral robusticity does not correlate directly with the level of logistical mobility, but is instead due to the summation of several diverse factors that place biomechanical loads on the hindlimb, particularly unevenness of the terrain. However, female femoral gracility seems to indicate that below a certain "threshold" of mobility, i.e., movement over the natural terrain, terrain conformation is no longer the main contributing factor to femoral robusticity. The femoral shape index I(x)/I(y) declines through time, particularly in males. This agrees with the expected mobility of the samples based on archaeological and historical data, providing further evidence on the reliability of this index in inferring terrestrial mobility.

Biomechanical approach to the reconstruction of activity patterns in Neolithic Western Liguria, Italy.

This paper investigates the changes in upper and lower limb robusticity and activity patterns that accompanied the transition to a Neolithic subsistence in western Liguria (Italy). Diaphyseal robusticity measures were obtained from cross-sectional geometric properties of the humerus and femur in a sample of 16 individuals (eight males and eight females) dated to about 6,000-5,500 BP. Comparisons with European Late Upper Paleolithics (LUP) indicate increased humeral robusticity in Neolithic Ligurian (NEOL) males, but not in females, with a significant reduction in right-left differences in both sexes. Sexual dimorphism in robusticity increases in upper and lower limb bones. Regarding the femur, while all female indicators of bending strength decrease steadily through time, values for NEOL males approach those of LUP. This suggests high, and unexpected, levels of mechanical stress for NEOL males, probably reflecting the effects of the mountainous terrain on lower limb remodeling. Comparisons between NEOL males and a small sample of LUP hunter-gatherers from the same area support this interpretation. In conclusion, cross-sectional geometry data indicate that the transition to Neolithic economies in western Liguria did not reduce functional requirements in males, and suggest a marked sexual division of labor involving a more symmetrical use of the upper limb, and different male-female levels of locomotory stress. When articulated with archaeological, faunal, paleopathological, and ethnographic evidence, these results support the hypothesis of repetitive, bimanual use of axes tied to pastoral activities in males, and of more sedentary tasks linked to agriculture in females.