The earliest ancient DNA data of modern humans from Europe dates to ~40 thousand 27 years ago1-4, but that from the Caucasus and the Near East to only ~14 thousand years 28 ago5,6, from populations who lived long after the Last Glacial Maximum (LGM) ~26.5-29 19 thousand years ago7. To address this imbalance and to better understand the 30 relationship of Europeans and Near Easterners, we report genome-wide data from two 31 ~26 thousand year old individuals from Dzudzuana Cave in Georgia in the Caucasus 32 from around the beginning of the LGM. Surprisingly, the Dzudzuana population was 33 more closely related to early agriculturalists from western Anatolia ~8 thousand years 34 ago8 than to the hunter-gatherers of the Caucasus from the same region of western 35 Georgia of ~13-10 thousand years ago5. Most of the Dzudzuana population’s ancestry 36 was deeply related to the post-glacial western European hunter-gatherers of the 37 ‘Villabruna cluster’3, but it also had ancestry from a lineage that had separated from 38 the great majority of non-African populations before they separated from each other, 39 proving that such ‘Basal Eurasians’6,9 were present in West Eurasia twice as early as 40 It is made available under a CC-BY-NC-ND 4.0 International license.

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bioRxiv preprint first posted online Sep. 21, 2018; doi: http://dx.doi.org/10.1101/423079. The copyright holder for this preprint

previously recorded5,6. We document major population turnover in the Near East after 41 the time of Dzudzuana, showing that the highly differentiated Holocene populations of 42 the region6 were formed by ‘Ancient North Eurasian’3,9,10 admixture into the Caucasus 43 and Iran and North African11,12 admixture into the Natufians of the Levant. We finally 44 show that the Dzudzuana population contributed the majority of the ancestry of post-45 Ice Age people in the Near East, North Africa, and even parts of Europe, thereby 46 becoming the largest single contributor of ancestry of all present-day West Eurasians.