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.

Grey wolf genomic history reveals a dual ancestry of dogs.

The grey wolf (Canis lupus) was the first species to give rise to a domestic population, and they remained widespread throughout the last Ice Age when many other large mammal species went extinct. Little is known, however, about the history and possible extinction of past wolf populations or when and where the wolf progenitors of the present-day dog lineage (Canis familiaris) lived1-8. Here we analysed 72 ancient wolf genomes spanning the last 100,000 years from Europe, Siberia and North America. We found that wolf populations were highly connected throughout the Late Pleistocene, with levels of differentiation an order of magnitude lower than they are today. This population connectivity allowed us to detect natural selection across the time series, including rapid fixation of mutations in the gene IFT88 40,000-30,000 years ago. We show that dogs are overall more closely related to ancient wolves from eastern Eurasia than to those from western Eurasia, suggesting a domestication process in the east. However, we also found that dogs in the Near East and Africa derive up to half of their ancestry from a distinct population related to modern southwest Eurasian wolves, reflecting either an independent domestication process or admixture from local wolves. None of the analysed ancient wolf genomes is a direct match for either of these dog ancestries, meaning that the exact progenitor populations remain to be located.

Stable isotope evidence of human diet in Mediterranean context during the Last Glacial Maximum.

The Final Gravettian in Europe overlapped with the cold and dry climatic event of Heinrich 2 (ca. 27-23.5 kyr cal BP), which caused the contraction of human distribution over refuge regions in the southern peninsulas of Europe. Here, we consider the human subsistence in the northeast Iberian Peninsula, where an extensive range of small to large prey was available. Four human remains from the Serinyà caves were investigated using the stable isotope ratios of carbon, nitrogen, and sulfur of bulk collagen (δ13Ccoll, δ15Ncoll, δ34Scoll) and of phenylalanine and glutamine amino acids (δ15NPhe, δ15NGlu). Direct AMS dating of the human and animal remains from the Final Gravettian levels of Mollet III, Reclau Viver, and Arbreda at Serinyà confirmed their chronological position from 27.5 to 22.6 kyr cal BP and the occurrence of four different human individuals. The δ13Ccoll and δ15Ncoll values showed a large contribution of terrestrial prey to the dietary protein of the individuals. The δ34Scoll values were consistent with a subsistence based on the local continental resources, without detectable contribution of marine resource. The δ15NPhe and δ15NGlu values confirm that freshwater resources were not a substantial component of the diet of the considered individuals. Contrast in the isotopic amounts in bulk collagen could be interpreted as the result of different proportions of terrestrial prey in human diet at Serinyà. Altogether, the isotopic investigation reveals the importance of terrestrial over aquatic resources in the subsistence of the studied Final Gravettian individuals from the Serinyà caves in northeastern Iberia during the Last Glacial Maximum. It would be consistent with a scenario of a productive enough terrestrial ecosystem to sustain hunter-gatherer subsistence in this refuge region.

The genomic history of southeastern Europe.

Farming was first introduced to Europe in the mid-seventh millennium bc, and was associated with migrants from Anatolia who settled in the southeast before spreading throughout Europe. Here, to understand the dynamics of this process, we analysed genome-wide ancient DNA data from 225 individuals who lived in southeastern Europe and surrounding regions between 12000 and 500 bc. We document a west-east cline of ancestry in indigenous hunter-gatherers and, in eastern Europe, the early stages in the formation of Bronze Age steppe ancestry. We show that the first farmers of northern and western Europe dispersed through southeastern Europe with limited hunter-gatherer admixture, but that some early groups in the southeast mixed extensively with hunter-gatherers without the sex-biased admixture that prevailed later in the north and west. We also show that southeastern Europe continued to be a nexus between east and west after the arrival of farmers, with intermittent genetic contact with steppe populations occurring up to 2,000 years earlier than the migrations from the steppe that ultimately replaced much of the population of northern Europe.

Determination of the dietary habits of a Magdalenian woman from Saint-Germain-la-Rivière in southwestern France using stable isotopes.

To obtain direct dietary information, carbon and nitrogen isotope ratios were measured from bone collagen acquired from the well-preserved skeleton of a Magdalenian woman from the site of Saint-Germain-la-Rivière in southwestern France. Comparison of delta13C and delta15N values of the human bone collagen to those of bone collagen from local herbivores and carnivores indicates that the woman's primary source of protein was the meat of large terrestrial herbivores. Application of a linear mixing model to the woman's isotopic signature indicates that (1) no significant marine-derived protein contributed to her average diet; (2) saiga antelope, which dominates the faunal remains at Saint-Germain-la-Rivière, was not the main source of terrestrial protein; and (3) her pattern of subsistence reflects a less opportunistic behavior than generally attributed to humans from this period. Dietary proportions of prey reflected by the number of identified specimens are revised using meat percentage estimates, which de-emphasize the importance of saiga antelope in human subsistence at Saint-Germain-la-Rivière during the middle Magdalenian.

Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: review and use of a multi-source mixing model.

The carbon and nitrogen isotopic abundances of the collagen extracted from the Saint-Césaire I Neanderthal have been used to infer the dietary behaviour of this specimen. A review of previously published Neanderthal collagen isotopic signatures with the addition of 3 new collagen isotopic signatures from specimens from Les Pradelles allows us to compare the dietary habits of 5 Neanderthal specimens from OIS 3 and one specimen from OIS 5c. This comparison points to a trophic position as top predator in an open environment, with little variation through time and space. In addition, a comparison of the Saint-Césaire I Neanderthal with contemporaneous hyaenas has been performed using a multi-source mixing model, modified from Phillips and Gregg (2003, Oecologia 127, 171). It appears that the isotopic differences between the Neanderthal specimen and hyaenas can be accounted for by much lower amounts of reindeer and much higher amounts of woolly rhinoceros and woolly mammoth in the dietary input of the Neanderthal specimen than in that of hyaenas, with relatively similar contributions of bovinae, large deer and horse for both predators, a conclusion consistent with the zooarchaeological data. The high proportion of very large herbivores, such as woolly rhinoceros and woolly mammoth, in Neanderthal's diet compare to that of the scavenging hyaenas suggests that Neanderthals could not acquire these prey through scavenging. They probably had to hunt for proboscideans and rhinoceros. Such a prey selection could result from a long lasting dietary tradition in Europe.