Dorothée G. Drucker

The genetic history of Ice Age Europe

Modern humans arrived in Europe ~45,000 years ago, but little is known about their genetic composition before the start of farming ~8,500 years ago. We analyze genome-wide data from 51 Eurasians from ~45,000-7,000 years ago. Over this time, the proportion of Neanderthal DNA decreased from 3–6% to around 2%, consistent with natural selection against Neanderthal variants in modern humans. Whereas the earliest modern humans in Europe did not contribute substantially to present-day Europeans, all individuals between ~37,000 and ~14,000 years ago descended from a single founder population which forms part of the ancestry of present-day Europeans. A ~35,000 year old individual from northwest Europe represents an early branch of this founder population which was then displaced across a broad region, before reappearing in southwest Europe during the Ice Age ~19,000 years ago. During the major warming period after ~14,000 years ago, a new genetic component related to present-day Near Easterners appears in Europe. These results document how population turnover and migration have been recurring themes of European pre-history.

Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe

How modern humans dispersed into Eurasia and Australasia, including the number of separate expansions and their timings, is highly debated [1, 2]. Two categories of models are proposed for the dispersal of non-Africans: (1) single dispersal, i.e., a single major diffusion of modern humans across Eurasia and Australasia [3-5]; and (2) multiple dispersal, i.e., additional earlier population expansions that may have contributed to the genetic diversity of some present-day humans outside of Africa [6-9]. Many variants of these models focus largely on Asia and Australasia, neglecting human dispersal into Europe, thus explaining only a subset of the entire colonization process outside of Africa [3-5, 8, 9]. The genetic diversity of the first modern humans who spread into Europe during the Late Pleistocene and the impact of subsequent climatic events on their demography are largely unknown. Here we analyze 55 complete human mitochondrial genomes (mtDNAs) of hunter-gatherers spanning ∼35,000 years of European prehistory. We unexpectedly find mtDNA lineage M in individuals prior to the Last Glacial Maximum (LGM). This lineage is absent in contemporary Europeans, although it is found at high frequency in modern Asians, Australasians, and Native Americans. Dating the most recent common ancestor of each of the modern non-African mtDNA clades reveals their single, late, and rapid dispersal less than 55,000 years ago. Demographic modeling not only indicates an LGM genetic bottleneck, but also provides surprising evidence of a major population turnover in Europe around 14,500 years ago during the Late Glacial, a period of climatic instability at the end of the Pleistocene.

Ancient DNA sequences point to a large loss of mitochondrial genetic diversity in the saiga antelope (Saiga tatarica) since the Pleistocene

Prior to the Holocene, the range of the saiga antelope (Saiga tatarica) spanned from France to the Northwest Territories of Canada. Although its distribution subsequently contracted to the steppes of Central Asia, historical records indicate that it remained extremely abundant until the end of the Soviet Union, after which its populations were reduced by over 95%. We have analysed the mitochondrial control region sequence variation of 27 ancient and 38 modern specimens, to assay how the species’ genetic diversity has changed since the Pleistocene. Phylogenetic analyses reveal the existence of two well‐supported, and clearly distinct, clades of saiga. The first, spanning a time range from >49 500 14C ybp to the present, comprises all the modern specimens and ancient samples from the Northern Urals, Middle Urals and Northeast Yakutia. The second clade is exclusive to the Northern Urals and includes samples dating from between 40 400 to 10 250 14C ybp. Current genetic diversity is much lower than that present during the Pleistocene, an observation that data modelling using serial coalescent indicates cannot be explained by genetic drift in a population of constant size. Approximate Bayesian Computation analyses show the observed data is more compatible with a drastic population size reduction (c. 66–77%) following either a demographic bottleneck in the course of the Holocene or late Pleistocene, or a geographic fragmentation (followed by local extinction of one subpopulation) at the Holocene/Pleistocene transition.

Influence of forage preferences and habitat use on 13C and 15N abundance in wild caribou (Rangifer tarandus caribou) and moose (Alces alces) from Canada.

Stable isotope composition (δ13C and δ15N) of moose (Alces alces) and caribou (Rangifer tarandus) hair from the boreal forest of Jacques-Cartier Park and Côte-Nord (Québec) and arctic tundra of Queen Maud Gulf and Southampton Island (Nunavut) was investigated as an indicator of dietary preferences and habitat use. Values of δ13Chair and δ15Nhair in moose were consistently lower compared to those of caribou. This is consistent with the depletion in 13C and 15N in the plants preferred by moose, essentially browse (shrub and tree leaves), compared to caribou forage, which included significant amounts of graminoids, lichen and fungi. The δ13Chair values of caribou differed between closed boreal forest and open-tundra ecosystems. This pattern followed that expected from the canopy effect observed in plant communities. Variation in δ15Nhair values of caribou was probably linked to the effect of different climatic conditions on plant communities. This study underlines the potential of isotopic analysis for studies on diet and habitat selection within a pure C3 plant environment.

Evidence for a 15N positive excursion in terrestrial foodwebs at the Middle to Upper Palaeolithic transition in south-western France: Implications for early modern human palaeodiet and palaeoenvironment

The Middle to Upper Palaeolithic transition around 35,000 years ago coincides with the replacement of Neanderthals by anatomically modern humans in Europe. Several hypotheses have been suggested to explain this replacement, one of them being the ability of anatomically modern humans to broaden their dietary spectrum beyond the large ungulate prey that Neanderthals consumed exclusively. This scenario is notably based on higher nitrogen-15 amounts in early Upper Palaeolithic anatomically modern human bone collagen compared with late Neanderthals. In this paper, we document a clear increase of nitrogen-15 in bone collagen of terrestrial herbivores during the early Aurignacian associated with anatomically modern humans compared with the stratigraphically older Châtelperronian and late Mousterian fauna associated with Neanderthals. Carnivores such as wolves also exhibit a significant increase in nitrogen-15, which is similar to that documented for early anatomically modern humans compared with Neanderthals in Europe. A shift in nitrogen-15 at the base of the terrestrial foodweb is responsible for such a pattern, with a preserved foodweb structure before and after the Middle to Upper Palaeolithic transition in south-western France. Such an isotopic shift in the terrestrial ecosystem may be due to an increase in aridity during the time of deposition of the early Aurignacian layers. If it occurred across Europe, such a shift in nitrogen-15 in terrestrial foodwebs would be enough to explain the observed isotopic trend between late Neanderthals and early anatomically modern humans, without any significant change in the diet composition at the Middle to Upper Palaeolithic transition.

European Bison as a Refugee Species? Evidence from Isotopic Data on Early Holocene Bison and Other Large Herbivores in Northern Europe

According to the refugee species concept, increasing replacement of open steppe by forest cover after the last glacial period and human pressure had together forced European bison (Bison bonasus)—the largest extant terrestrial mammal of Europe—into forests as a refuge habitat. The consequent decreased fitness and population density led to the gradual extinction of the species. Understanding the pre-refugee ecology of the species may help its conservation management and ensure its long time survival. In view of this, we investigated the abundance of stable isotopes (δ13C and δ15N) in radiocarbon dated skeletal remains of European bison and other large herbivores—aurochs (Bos primigenius), moose (Alces alces), and reindeer (Rangifer tarandus)—from the Early Holocene of northern Europe to reconstruct their dietary habits and pattern of habitat use in conditions of low human influence. Carbon and nitrogen isotopic compositions in collagen of the ungulate species in northern central Europe during the Early Holocene showed significant differences in the habitat use and the diet of these herbivores. The values of the δ13C and δ15N isotopes reflected the use of open habitats by bison, with their diet intermediate between that of aurochs (grazer) and of moose (browser). Our results show that, despite the partial overlap in carbon and nitrogen isotopic values of some species, Early Holocene large ungulates avoided competition by selection of different habitats or different food sources within similar environments. Although Early Holocene bison and Late Pleistocene steppe bison utilized open habitats, their diets were significantly different, as reflected by their δ15N values. Additional isotopic analyses show that modern populations of European bison utilize much more forested habitats than Early Holocene bison, which supports the refugee status of the species.

The Oldest Anatomically Modern Humans from Far Southeast Europe: Direct Dating, Culture and Behavior

Background Anatomically Modern Humans (AMHs) are known to have spread across Europe during the period coinciding with the Middle to Upper Paleolithic transition. Whereas their dispersal into Western Europe is relatively well established, evidence of an early settlement of Eastern Europe by modern humans are comparatively scarce. Methodology/Principal Finding Based on a multidisciplinary approach for the study of human and faunal remains, we describe here the oldest AMH remains from the extreme southeast Europe, in conjunction with their associated cultural and paleoecological background. We applied taxonomy, paleoecology, and taphonomy combined with geomorphology, stratigraphy, archeology and radiocarbon dating. More than 160 human bone remains have been discovered. They originate from a well documented Upper Paleolithic archeological layer (Gravettian cultural tradition) from the site of Buran-Kaya III located in Crimea (Ukraine). The combination of non-metric dental traits and the morphology of the occipital bones allow us to attribute the human remains to Anatomically Modern Humans. A set of human and faunal remains from this layer has been radiocarbon dated by Accelerator Mass Spectrometry. The direct-dating results of human bone establish a secure presence of AMHs at 31,900+240/−220 BP in this region. They are the oldest direct evidence of the presence of AMHs in a well documented archeological context. Based on taphonomical observations (cut marks and distribution of skeletal elements), they represent the oldest Upper Paleolithic modern humans from Eastern Europe, showing post-mortem treatment of the dead as well. Conclusion/Significance These findings are essential for the debate on the spread of modern humans in Europe during the Upper Paleolithic, as well as their cultural behaviors.

Investigation of equid paleodiet from Schöningen 13 II-4 through dental wear and isotopic analyses: Archaeological implications

The paleodietary traits of the equid population from Schöningen 13 II-4 were investigated through tooth mesowear and microwear analyses, as well as stable isotopic analyses. The mesowear pattern observed on the upper teeth indicates a low abrasion diet with a significant amount of browse in the diet of the horses. The tooth microwear analysis and the isotopic data confirm that the horses from Schöningen 13 II-4 were mixed feeders, like many populations from other Pleistocene localities in Northern and Eastern Europe. Microwear also provides information on seasonal changes in the diet of the horses and offers the possibility to test hypotheses about the presence of one or several horse populations. Our analysis determined that the assemblage of horse remains from Schöningen 13 II-4 resulted from multiple accumulation events, which took place at different periods of time.

Ecological niche of Neanderthals from Spy Cave revealed by nitrogen isotopes of individual amino acids in collagen

This study provides a refined view on the diet and ecological niche of Neanderthals. The traditional view is that Neanderthals obtained most of their dietary protein from terrestrial animals, especially from large herbivores that roamed the open landscapes. Evidence based on the conventional carbon and nitrogen isotopic composition of bulk collagen has supported this view, although recent findings based on plant remains in the tooth calculus, microwear analyses, and small game and marine animal remains from archaeological sites have raised some questions regarding this assumption. However, the lack of a protein source other than meat in the Neanderthal diet may be due to methodological difficulties in defining the isotopic composition of plants. Based on the nitrogen isotopic composition of glutamic acid and phenylalanine in collagen for Neanderthals from Spy Cave (Belgium), we show that i) there was an inter-individual dietary heterogeneity even within one archaeological site that has not been evident in bulk collagen isotopic compositions, ii) they occupied an ecological niche different from those of hyenas, and iii) they could rely on plants for up to ∼20% of their protein source. These results are consistent with the evidence found of plant consumption by the Spy Neanderthals, suggesting a broader subsistence strategy than previously considered.

Carbon and nitrogen stable isotopes of well-preserved Middle Pleistocene bone collagen from Schoningen (Germany) and their paleoecological implications

Carbon and nitrogen stable isotopes in bone collagen can provide valuable information about the diet and habitat of mammal species. However, bone collagen degrades in normal circumstances very rapidly, and isotope analyses are therefore usually restricted to fossil material with a Late Pleistocene or Holocene age. The Middle Pleistocene site of Schöningen, dated to around 300,000 years ago, yielded bones and teeth with an exceptionally good state of collagen preservation. This allowed us to measure reliable biogenic carbon and nitrogen stable isotope ratios for different herbivorous taxa from the families Elephantidae, Rhinocerotidae, Equidae, Cervidae, and Bovidae. The results provide insights regarding the paleoenvironmental setting in which Middle Pleistocene hominins operated. The vegetation consumed by the herbivores from the famous spear horizon originates from open environments. During the climatic Reinsdorf Interglacial optimum, the landscape seems to have been relatively open as well, but certainly included parts that were forested. The results also indicate some niche partitioning; different herbivore species used different plant resources. For instance, the horses seem to have been predominantly browsers, while the straight-tusked elephants were feeding chiefly on grass.