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Figure 1 : Genetic structure of ancient Europe.


We extend the scope of European palaeogenomics by sequencing the genomes of Late Upper Palaeolithic (13,300 years old, 1.4-fold coverage) and Mesolithic (9,700 years old, 15.4-fold) males from western Georgia in the Caucasus and a Late Upper Palaeolithic (13,700 years old, 9.5-fold) male from Switzerland. While we detect Late Palaeolithic–Mesolithic genomic continuity in both regions, we find that Caucasus hunter-gatherers (CHG) belong to a distinct ancient clade that split from western hunter-gatherers 45 kya, shortly after the expansion of anatomically modern humans into Europe and from the ancestors of Neolithic farmers 25  kya, around the Last Glacial Maximum. CHG genomes significantly contributed to the Yamnaya steppe herders who migrated into Europe 3,000 BC, supporting a formative Caucasus influence on this important Early Bronze age culture. CHG left their imprint on modern populations from the Caucasus and also central and south Asia possibly marking the arrival of Indo-Aryan languages.


Ancient genomes from Eurasia have revealed three ancestral populations that contributed to contemporary Europeans in varying degrees1. Mesolithic individuals, sampled from Spain all the way to Hungary1,2,3, belong to a relatively homogenous group, termed western hunter-gatherers (WHG). The expansion of early farmers (EF) out of the Levant during the Neolithic transition led to major changes in the European gene pool, with almost complete replacement in the south and increased mixing with local WHG further north1,2,3,4,5. Finally, a later wave originating with the Early Bronze Age Yamnaya from the Pontic steppe, carrying partial ancestry from ancient North Eurasians (ANE) and ancestry from a second, undetermined source, arrived from the east, profoundly changing populations and leaving a cline of admixture in Eastern and Central Europe1,3,6. This view, which was initially based on a handful of genomes, was recently confirmed by extensive surveys of Eurasian samples from the Holocene5,7.

Here, we extend our view of the genetic makeup of early Europeans by both looking further back in time and sampling from the crossroads between the European and Asian continents. We sequenced a Late Upper Palaeolithic (‘Satsurblia’ from Satsurblia cave, 1.4-fold coverage) and a Mesolithic genome (‘Kotias’ from Kotias Klde cave, 15.4-fold) from Western Georgia, at the very eastern boundary of Europe. We term these two individuals Caucasus hunter-gatherers (CHG). To extend our overview of WHG to a time depth similar to the one available for our samples from the Caucasus, we also sequenced a western European Late Upper Palaeolithic genome, ‘Bichon’ (9.5-fold) from Grotte du Bichon, Switzerland. These new genomes, together with already published data, provide us with a much-improved geographic and temporal coverage of genetic diversity across Europe after the Last Glacial Maximum (LGM)8. We show that CHG belong to a new, distinct ancient clade that split from WHG 45 kya and from Neolithic farmer ancestors 25  kya. This clade represents the previously undetermined source of ancestry to the Yamnaya, and contributed directly to modern populations from the Caucasus all the way to Central Asia.

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