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However, this interaction zone predates any direct influence of Yamnaya groups in Europe or the succeeding formation of the Corded Ware 46 , 47 and its persistence opens the possibility of subtle gene-flow from farmers at the eastern border of arable lands into the steppe, several centuries before the massive range expansions of pastoralist groups that reached Central Europe in the mid-3 rd millennium BCE 18 , A surprising discovery was that Steppe Maykop individuals from the eastern desert steppes harboured a distinctive ancestry component that relates them to Upper Palaeolithic Siberians AG3, MA1 and Native Americans.

The additional affinity to East Asians suggests that this ancestry is not derived directly from ANE but from a yet-to-be-identified ancestral population in north-central Eurasia with a wide distribution between the Caucasus, the Ural Mountains and the Pacific coast 20 , of which we have discovered the so far southwestern-most and also youngest genetic representatives. The insight that the Caucasus mountains served as a corridor for the spread of CHG ancestry north but also for subtle later gene-flow from the south allows speculations on the postulated homelands of Proto-Indo-European PIE languages and documented gene-flows that could have carried a consecutive spread of both across West Eurasia 15 , This also opens up the possibility of a homeland of PIE south of the Caucasus, and could offer a parsimonious explanation for an early branching off of Anatolian languages, as shown on many PIE tree topologies 50 , 51 , 52 , Geographically conceivable are also Armenian and Greek, for which genetic data support an eastern influence from Anatolia or the southern Caucasus 10 , 54 , and an Indo-Iranian offshoot to the east.

Samples from archaeological human remains were collected and exported under a collaborative research agreement between the Max-Planck Institute for the Science of Human History, the German Archaeological Institute and the Lomonosov Moscow State University and Anuchin Research Institute and Museum of Anthropology permission no. We extracted DNA and prepared next-generation sequencing libraries from samples in two dedicated ancient DNA laboratories at Jena and Boston, following established protocols for DNA extraction and library preparation 55 , Teeth were sandblasted to remove the outer surface and then ground to fine powder using a mixer mill Retsch, Germany.

We also sampled the dense parts of petrous bones by cutting out a bone wedge around the region of the cochlea, which—after surface removal—was also ground to fine bone powder. The lysis step included the addition of extraction buffer, containing 0. Blank controls were processed in parallel at a ratio of We then added 0. Blunt-end-repair of the DNA fragments was carried out by adding 0. Illumina adaptors 0. The final fill-in step included 1X isothermal buffer, 0.

For those libraries passing quality threshholds, we carried out in-solution enrichment K capture 22 for a targeted set of 1,, SNPs as well as mitochondrial genome capture, and then sequenced on for 76bp either single or paired-end. We report, but have not analyzed, data from individuals that had less than 30, SNP hits on the K set.

We removed individuals with evidence of contamination based on heterozygosity in the mtDNA genome data, a high rate of heterozygosity on the X chromosome despite being male estimated with ANGSD 60 , or an atypical ratio of the reads mapped to X versus Y chromosomes.

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We also included newly genotyped populations from the Caucasus and Asia, described in detail in Jeong et al. DG, Papuan. DG, Onge. SG, Karitiana. For some analyses, we used an extended set of outgroup populations, including some of the following additional ancient populations to constrain standard errors: WHG, EHG, and Levant Neolithic.

We estimated the time depth of selected admixture events using the linkage disequilibrium LD -based admixture inference implemented in ALDER 32 , assuming a generation time of 28 years DG as an outgroup. We explored models that jointly explain the population splits and gene flow in the Greater Caucasus region by computing f 2 -, f 3 - and f 4 - statistics measuring allele sharing among pairs, triples, and quadruples of populations and evaluating fits based on the maximum Z -score comparing predicted and observed values of these statistics.

We determined the sex of the newly reported samples in this study by counting the number of reads overlapping with the targets of k capture reagent We extracted the reads of high base and mapping quality samtools depth -q30 -Q37 using samtools v1. We used Geneious R8.


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In addition, we used the software contamMix 1. We determined Y chromosomal haplogroups by identifying the most derived allele upstream and the most ancestral allele downstream in the phylogenetic tree in the ISOGG version We used outgroup- f 3 statistics and the methods lcMLkin 69 and READ 70 to determine genetic kinship between individuals. Adler, D. Science , — Pinhasi, R. New chronology for the Middle Palaeolithic of the southern Caucasus suggests early demise of Neanderthals in this region. Lordkipanidze, D. A complete skull from Dmanisi, Georgia, and the evolutionary biology of early Homo.

Helwing, B. The Kura projects: new research on the later prehistory of the southern Caucasus. Kohl, P. The prehistory of the caucasus: internal developments and external interactions.

Introduction

Cambridge University Press, Cambridge, Stein, G. The development of indigenous social complexity in late Chalcolithic Upper Mesopotamia in the 5th-4th Millennia BC—an initial assessment. Origini 34 , — The Making of Bronze Age Eurasia.

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Govedarica, B. Philipp von Zabern, Mainz am Rhein, Lazaridis, I. Genomic insights into the origin of farming in the ancient Near East. Nature , — Mathieson, I.

3, Decorative Patterns of the Ancient World (eBook)

The genomic history of southeastern Europe. Benecke, N.

Finding the Woolly Sheep. Meta-analyses of archaeozoological data from Southwest-Asia and Southeast-Europe. Reinhold S. Maran, J, Stockhammer, P. Jones, E. Upper Palaeolithic genomes reveal deep roots of modern Eurasians. Anthony, D. The secondary products revolution, horse-riding, and mounted warfare. World Prehist. Frachetti, M. Multiregional emergence of mobile pastoralism and nonuniform institutional complexity across Eurasia. Librado, P.

The evolutionary origin and genetic makeup of domestic horses. Genetics , — Haak, W. Massive migration from the steppe was a source for Indo-European languages in Europe. Allentoft, M. Population genomics of Bronze Age Eurasia. Narasimhan, V. The genomic formation of south and central Asia. Ancient human genomes suggest three ancestral populations for present-day Europeans. Genome-wide patterns of selection in ancient Eurasians. Broushaki, F. Early Neolithic genomes from the eastern Fertile Crescent.

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A brief history of Western culture

The caucasus as an asymmetric semipermeable barrier to ancient human migrations. Balanovsky, O. Parallel evolution of genes and languages in the Caucasus region. Schonberg, A. High-throughput sequencing of complete human mtDNA genomes from the Caucasus and West Asia: high diversity and demographic inferences. Fu, Q. An early modern human from Romania with a recent Neanderthal ancestor.


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Patterson, N. Ancient admixture in human history. Alexander, D. Fast model-based estimation of ancestry in unrelated individuals.