The formation of human populations in South and Central Asia

RATIONALE To elucidate the extent to which the major cultural transformations of farming, pastoralism, and shifts in the distribution of languages in Eurasia were accompanied by movement of people, we report genome-wide ancient DNAdata from 523 individuals spanning the last 8000 years, mostly from Central Asia and northernmost South Asia. RESULTS The movement of people following the advent of farming resulted in genetic gradients across Eurasia that can be modeled as mixtures of seven deeply divergent populations. A key gradient formed in southwestern Asia beginning in the Neolithic and continuing into the Bronze Age, with more Anatolian farmer–related ancestry in the west and more Iranian farmer–related ancestry in the east. This cline extended to the desert oases of Central Asia and was the primary source of ancestry in peoples of the Bronze AgeBactria Margiana Archaeological Complex (BMAC). This supports the idea that the archaeologically documented dispersal of domesticates was accompanied by the spread of people from multiple centers of domestication. The main population of the BMAC carried no ancestry from Steppepastoralists and did not contribute substantially to later South Asians. However, Steppepastoralist ancestry appeared in outlier individuals at BMAC sites by the turn of the second millennium BCE around the same time as it appeared on the southern Steppe. Using data from ancient individuals from the Swat Valley of northernmost South Asia, we show that Steppeancestry then integrated further south in the first half of the second millennium BCE, contributing up to 30% of the ancestry of modern groups in South Asia. The Steppeancestry in South Asia has the same profile as that in Bronze AgeEastern Europe, tracking a movement of people that affected both regions and that likely spread the unique features shared between Indo-Iranian and Balto- Slavic languages. The primary ancestral population of modern South Asians is a mixture of people related to early Holocene populations of Iran and South Asia that we detect in outlier individuals from two sites in cultural contact with the IndusValley Civilization (IVC), making it plausible that it was characteristic of the IVC. After the IVC’s decline, this population mixed with northwestern groups with Steppeancestry to form the “Ancestral North Indians” (ANI) and also mixed with southeastern groups to form the “Ancestral South Indians” (ASI), whose direct descendants today live in tribal groups in southern India. Mixtures of these two post-IVC groups—the ANI and ASI—drive the main gradient of genetic variation in South Asia today. CONCLUSION Earlier work recorded massive population movement from the Eurasian Steppeinto Europe early in the third millennium BCE, likely spreading Indo-European languages. We reveal a parallel series of events leading to the spread of Steppeancestry to South Asia, thereby documenting movements of people that were likely conduits for the spread of Indo-European languages.