Migrations and mixing

The proportion of Neanderthal-derived DNA in present-day people outside Africa is estimated at 1.5-2.1%.

Prüfer K., Racimo F., Patterson N. et al. (2014). The complete genome sequence of a Neanderthal from the Altai Mountains. Nature 505, 43-49. doi:10.1038/nature12886

Bergström et al. (2020) cite the consensus view that modern humans expanded out of Africa and the Near East 50,000-70,000 years ago.

Bergström A., McCarthy S. A., Hui R. et al. (2020). Insights into human genetic variation and population history from 929 diverse genomes. Science 367(6484), eaay5012. doi:10.1126/science.aay5012

The two infants from the Upward Sun River site (Alaska) are dated to around 11,500 years ago.

Moreno-Mayar J. V., Potter B. A., Vinner L. et al. (2018). Terminal Pleistocene Alaskan genome reveals first founding population of Native Americans. Nature 553, 203-207. doi:10.1038/nature25173

The Upward Sun River individual USR1 represents a distinct Ancient Beringian population.

Moreno-Mayar J. V., Potter B. A., Vinner L. et al. (2018). Terminal Pleistocene Alaskan genome reveals first founding population of Native Americans. Nature 553, 203-207. doi:10.1038/nature25173

A single founding population of Native Americans split from East Asians around 36 ± 1.5 thousand years ago, with gene flow continuing until around 25 ± 1.1 thousand years ago; Ancient Beringians branched off around 22-18.1 thousand years ago.

Moreno-Mayar J. V., Potter B. A., Vinner L. et al. (2018). Terminal Pleistocene Alaskan genome reveals first founding population of Native Americans. Nature 553, 203-207. doi:10.1038/nature25173

Human footprints at White Sands National Park (New Mexico) are bracketed by seed layers with calibrated ages between about 23,000 and 21,000 years.

Bennett M. R., Bustos D., Pigati J. S. et al. (2021). Evidence of humans in North America during the Last Glacial Maximum. Science 373(6562), 1528-1531. doi:10.1126/science.abg7586

Northeastern Siberia was initially peopled by a previously unknown Palaeolithic population, the 'Ancient North Siberians', distantly related to early West Eurasian hunter-gatherers; the study's 34 ancient genomes date to between 31,000 and 600 years ago.

Sikora M., Pitulko V. V., Sousa V. C. et al. (2019). The population history of northeastern Siberia since the Pleistocene. Nature 570, 182-188. doi:10.1038/s41586-019-1279-z

The genetic makeup of northern Europe was shaped by migrations from Siberia that began at least 3,500 years ago; this Siberian ancestry is found particularly in populations speaking Uralic languages today.

Lamnidis T. C., Majander K., Jeong C. et al. (2018). Ancient Fennoscandian genomes reveal origin and spread of Siberian ancestry in Europe. Nature Communications 9, 5018. doi:10.1038/s41467-018-07483-5

The 3,500-year-old individuals from Bolshoy Oleni Ostrov (Murmansk Region) carry the highest proportion of Siberian-related ancestry in the region.

Lamnidis T. C., Majander K., Jeong C. et al. (2018). Ancient Fennoscandian genomes reveal origin and spread of Siberian ancestry in Europe. Nature Communications 9, 5018. doi:10.1038/s41467-018-07483-5

The Jomon diverged deeply from continental populations 20,000 to 15,000 years ago and maintained a small effective population size of about 1,000 over several millennia.

Cooke N. P., Mattiangeli V., Cassidy L. M. et al. (2021). Ancient genomics reveals tripartite origins of Japanese populations. Science Advances 7(38), eabh2419. doi:10.1126/sciadv.abh2419

Rice cultivation was introduced by people with Northeast Asian ancestry (Yayoi period), and a later influx of East Asian ancestry occurred during the imperial Kofun period.

Cooke N. P., Mattiangeli V., Cassidy L. M. et al. (2021). Ancient genomics reveals tripartite origins of Japanese populations. Science Advances 7(38), eabh2419. doi:10.1126/sciadv.abh2419

Genome-wide ancient DNA from 44 ancient Near Easterners living between about 12,000 and 1,400 BC shows that the first farmers of the southern Levant (Israel and Jordan) and the Zagros Mountains (Iran) were strongly genetically differentiated.

Lazaridis I., Nadel D., Rollefson G. et al. (2016). Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419-424. doi:10.1038/nature19310

Farmers related to those of Anatolia spread westward into Europe, those related to the Levant spread southward into East Africa, and those related to Iran spread northward into the Eurasian steppe.

Lazaridis I., Nadel D., Rollefson G. et al. (2016). Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419-424. doi:10.1038/nature19310

Most present-day Europeans derive from at least three highly differentiated populations: West European hunter-gatherers, Ancient North Eurasians related to Upper Palaeolithic Siberians, and Early European Farmers of mainly Near Eastern origin.

Lazaridis I., Patterson N., Mittnik A. et al. (2014). Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513, 409-413. doi:10.1038/nature13673

Neolithic Anatolians were members of the population that was the source of Europe's first farmers.

Mathieson I., Lazaridis I., Rohland N. et al. (2015). Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528, 499-503. doi:10.1038/nature16152

Farming arrived in Europe around 8,500 years ago.

Mathieson I., Lazaridis I., Rohland N. et al. (2015). Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528, 499-503. doi:10.1038/nature16152

Language phylogenies place the Austronesian origin in Taiwan approximately 5,230 years ago.

Gray R. D., Drummond A. J., Greenhill S. J. (2009). Language phylogenies reveal expansion pulses and pauses in Pacific settlement. Science 323(5913), 479-483. doi:10.1126/science.1166858

Ancient individuals from Vanuatu (about 3,100-2,700 years before present) and Tonga (about 2,700-2,300 years before present) had little to no Papuan ancestry.

Skoglund P., Posth C., Sirak K. et al. (2016). Genomic insights into the peopling of the Southwest Pacific. Nature 538, 510-513. doi:10.1038/nature19844

Western and Eastern Europe came into contact about 4,500 years ago; the Late Neolithic Corded Ware people from Germany traced about 75% of their ancestry to the Yamnaya.

Haak W., Lazaridis I., Patterson N. et al. (2015). Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522, 207-211. doi:10.1038/nature14317

The results support a steppe origin of at least some of the Indo-European languages of Europe.

Haak W., Lazaridis I., Patterson N. et al. (2015). Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522, 207-211. doi:10.1038/nature14317

Low-coverage genomes of 101 ancient humans show that the Bronze Age of Eurasia (around 3000-1000 BC) was a highly dynamic period involving large-scale population migrations and replacements.

Allentoft M. E., Sikora M., Sjögren K.-G. et al. (2015). Population genomics of Bronze Age Eurasia. Nature 522, 167-172. doi:10.1038/nature14507

Bantu-speaking peoples expanded about 4,000 to 5,000 years ago from western central Africa; the Bantu heartland is the Nigeria/Cameroon frontier.

Patin E., Lopez M., Grollemund R. et al. (2017). Dispersals and genetic adaptation of Bantu-speaking populations in Africa and North America. Science 356(6337), 543-546. doi:10.1126/science.aal1988

Early Bantu speakers first moved southward through the equatorial rainforest before spreading toward eastern and southern Africa.

Patin E., Lopez M., Grollemund R. et al. (2017). Dispersals and genetic adaptation of Bantu-speaking populations in Africa and North America. Science 356(6337), 543-546. doi:10.1126/science.aal1988

Based on 523 ancient humans: descendants of Steppe pastoralists, starting around 4,000 years ago, spread via Central Asia to form one of the main ancestral populations of South Asians.

Narasimhan V. M., Patterson N., Moorjani P. et al. (2019). The formation of human populations in South and Central Asia. Science 365(6457), eaat7487. doi:10.1126/science.aat7487

Steppe ancestry integrated further south, into South Asia, in the first half of the second millennium BCE.

Narasimhan V. M., Patterson N., Moorjani P. et al. (2019). The formation of human populations in South and Central Asia. Science 365(6457), eaat7487. doi:10.1126/science.aat7487

The study sequenced the genomes of 137 ancient humans (about 1× average coverage), covering a period of 4,000 years.

Damgaard P. de B., Marchi N., Rasmussen S. et al. (2018). 137 ancient human genomes from across the Eurasian steppes. Nature 557, 369-374. doi:10.1038/s41586-018-0094-2

Scythians admixed with eastern steppe nomads who formed the Xiongnu confederations; their descendants formed the Hun traditions in the fourth-fifth century AD.

Damgaard P. de B., Marchi N., Rasmussen S. et al. (2018). 137 ancient human genomes from across the Eurasian steppes. Nature 557, 369-374. doi:10.1038/s41586-018-0094-2

These changes transformed the Eurasian steppes from being inhabited by Indo-European speakers of largely West Eurasian ancestry to the mostly Turkic-speaking groups of the present day, who are primarily of East Asian ancestry.

Damgaard P. de B., Marchi N., Rasmussen S. et al. (2018). 137 ancient human genomes from across the Eurasian steppes. Nature 557, 369-374. doi:10.1038/s41586-018-0094-2

Genome-wide data from 555 ancient individuals, including 359 from Slavic contexts, demonstrate large-scale population movement from Eastern Europe during the sixth to eighth centuries CE.

Gretzinger J., Biermann F., Mager H. et al. (2025). Ancient DNA connects large-scale migration with the spread of Slavs. Nature 646, 384-393. doi:10.1038/s41586-025-09437-6

This movement replaced more than 80% of the local gene pool in Eastern Germany, Poland and Croatia, with substantial regional heterogeneity and no sex-biased admixture.

Gretzinger J., Biermann F., Mager H. et al. (2025). Ancient DNA connects large-scale migration with the spread of Slavs. Nature 646, 384-393. doi:10.1038/s41586-025-09437-6