We generated genome-wide data from 69 Europeans who lived between 8,000-3,000 years ago by enriching ancient DNA libraries for a target set of almost four hundred thousand polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies1–8 and to obtain new insights about the past. We show that the populations of western and far eastern Europe followed opposite trajectories between 8,000-5,000 years ago. At the beginning of the Neolithic period in Europe, ~8,000-7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary, and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ~24,000 year old Siberian6. By ~6,000-5,000 years ago, a resurgence of hunter-gatherer ancestry had occurred throughout much of Europe, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ~4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ~3/4 of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ~3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for the theory of a steppe origin9 of at least some of the Indo-European languages of Europe.

Bell Beaker pottery spread across western and central Europe beginning around 2750 BCE before disappearing between 2200–1800 BCE. The mechanism of its expansion is a topic of long-standing debate, with support for both cultural diffusion and human migration. We present new genome-wide ancient DNA data from 170 Neolithic, Copper Age and Bronze Age Europeans, including 100 Beaker-associated individuals. In contrast to the Corded Ware Complex, which has previously been identified as arriving in central Europe following migration from the east, we observe limited genetic affinity between Iberian and central European Beaker Complex-associated individuals, and thus exclude migration as a significant mechanism of spread between these two regions. However, human migration did have an important role in the further dissemination of the Beaker Complex, which we document most clearly in Britain using data from 80 newly reported individuals dating to 3900–1200 BCE. British Neolithic farmers were genetically similar to contemporary populations in continental Europe and in particular to Neolithic Iberians, suggesting that a portion of the farmer ancestry in Britain came from the Mediterranean rather than the Danubian route of farming expansion. Beginning with the Beaker period, and continuing through the Bronze Age, all British individuals harboured high proportions of Steppe ancestry and were genetically closely related to Beaker-associated individuals from the Lower Rhine area. We use these observations to show that the spread of the Beaker Complex to Britain was mediated by migration from the continent that replaced >90% of Britain’s Neolithic gene pool within a few hundred years, continuing the process that brought Steppe ancestry into central and northern Europe 400 years earlier.

Farming was first introduced to southeastern Europe in the mid-7th millennium BCE – brought by migrants from Anatolia who settled in the region before spreading throughout Europe. To clarify the dynamics of the interaction between the first farmers and indigenous hunter-gatherers where they first met, we analyze genome-wide ancient DNA data from 223 individuals who lived in southeastern Europe and surrounding regions between 12,000 and 500 BCE. We document previously uncharacterized genetic structure, showing a West-East cline of ancestry in hunter-gatherers, and show that some Aegean farmers had ancestry from a different lineage than the northwestern Anatolian lineage that formed the overwhelming ancestry of other European farmers. We show that the first farmers of northern and western Europe passed through southeastern Europe with limited admixture with local hunter-gatherers, but that some groups mixed extensively, with relatively sex-balanced admixture compared to the male-biased hunter-gatherer admixture that prevailed later in the North and West. Southeastern Europe continued to be a nexus between East and West after farming arrived, with intermittent genetic contact from the Steppe up to 2,000 years before the migration that replaced much of northern Europe’s population.

Domestication of horses fundamentally transformed long-range mobility and warfare1. However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling2–4 at Botai, Central Asia around 3500 bc3. Other longstanding candidate regions for horse domestication, such as Iberia5 and Anatolia6, have also recently been challenged. Thus, the genetic, geographic and temporal origins of modern domestic horses have remained unknown. Here we pinpoint the Western Eurasian steppes, especially the lower Volga-Don region, as the homeland of modern domestic horses. Furthermore, we map the population changes accompanying domestication from 273 ancient horse genomes. This reveals that modern domestic horses ultimately replaced almost all other local populations as they expanded rapidly across Eurasia from about 2000 bc, synchronously with equestrian material culture, including Sintashta spoke-wheeled chariots. We find that equestrianism involved strong selection for critical locomotor and behavioural adaptations at the GSDMC and ZFPM1 genes. Our results reject the commonly held association7 between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europe around 3000 bc8,9 driving the spread of Indo-European languages10. This contrasts with the scenario in Asia where Indo-Iranian languages, chariots and horses spread together, following the early second millennium bc Sintashta culture11,12.

The cemetery is located in the south-west of Pottenbrunn, on plot "Steinfeld" (15°41´05"/48°13´55"). Discovered in 1930, it had already yielded objects dating to the early La Tène period. In 1981, road construction revealed further finds which initiated rescue excavations by the Bundesdenkmalamt (State Office for Protection of Historical Monuments) under the guidance of J.-W. Neugebauer (Ramsl 2002a(Ramsl , 13) in 1981(Ramsl and 1982. A total of 42 graves with 45 burials (single and double inhumations, and cremations) have been documented. Some burials were severely disturbed (by ancient activities such as grave robbing and/or contemporary construction work), and some were set within fenced enclosures ("Grabgärten"). Three (of 22) samples of charcoal and bone fragments taken by Peter Stadler (Department of Prehistory, Natural History Museum Vienna) in the course of the FWFproject "Absolute Chronology for Early Civilisations in Austria and Central Europe" returned AMS dates of 410-200 cal BCE (grave 520), 550-200 cal BCE (grave 565) and 380-350 cal BCE (grave 1005) (Ramsl 2002b, 359). The cremation burials were not included in the initial osteological analysis, but 31 inhumed individuals were studied (Gerold 2002). Petrous bones from three of these were successfully analyzed for aDNA. Sample I11699 (female) derived from an individual (inv. no. 26.238) aged c. 20 years in grave 89 which, despite disturbance in antiquity, was accompanied by fibulae and ceramic vessels. Sample I11701 (male) derived from an individual (inv. no. 26.249) aged c. 18 years in grave 570, which also included shears, fibulae, and ceramic vessels. Evidence for bone porosity in the mandible and maxilla suggest possible Vitamin C deficiency, while enamel hypoplasia points to malnutrition or illness during childhood. Sample I11708 (female) derived from an individual (inv.no. 26.250) aged c. 25-35 years in grave 574/2, who was richly adorned with fibulae, bronze, iron and silver-rings, an amber ring, a bracelet, a glass bead, and a worked bone artefact.

Ancient DNA studies have established that European Neolithic populations were descended from Anatolian migrants who received a limited amount of admixture from resident hunter-gatherers. Many open questions remain, however, about the spatial and temporal dynamics of population interactions and admixture during the Neolithic period. Using the highest-resolution genome-wide ancient DNA data set assembled to date—a total of 177 samples, 127 newly reported here, from the Neolithic and Chalcolithic of Hungary (6000–2900 BCE, n = 98), Germany (5500–3000 BCE, n = 42), and Spain (5500–2200 BCE, n = 37)—we investigate the population dynamics of Neolithization across Europe. We find that genetic diversity was shaped predominantly by local processes, with varied sources and proportions of hunter-gatherer ances try among the three regions and through time. Admixture between groups with different ancestry profiles was pervasive and resulted in observable population transformation across almost all cultural transitions. Our results shed new light on the ways that gene flow reshaped European populations throughout the Neolithic period and demonstrate the potential of time-series-based sampling and modeling approaches to elucidate multiple dimensions of historical population interactions.

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Two sixth millennium cal BC cultural formations from the Carpathian Basin are discussed: the Linearbandkeramik (LBK) in Transdanubia, and the Szakálhát culture from the Great Hungarian Plain. Our aim was to address genetic connections between these cultural units and how they were related to the populations of the central European Neolithic. Mitochondrial data from 33 specimens from the Szakálhát culture were compared with 39 samples from the LBK and analysed in the light of previously published ancient DNA studies. The maternal gene pool of the LBK in Transdanubia and the Szakálhát culture are highly similar to each other. Furthermore, both groups show a remarkable affinity to the LBK and the subsequent fifth–fourth millennia cal BC cultures in central Europe. On the other hand, these Neolithic cultural formations can be clearly distinguished from Mesolithic pan-European hunter-gatherer data as well as from published Iberian Neolithic and central European Late Neolithic cultures.