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The Formation of Human Populations in South and Central Asia Vagheesh M

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The Formation of Human Populations in South and Central Asia Vagheesh M RESEARCH ◥ early Holocene populations of Iran and South RESEARCH ARTICLE SUMMARY Asia that we detect in outlier individuals from two sites in cultural contact with the Indus Valley Civilization (IVC), making it plausible HUMAN EVOLUTION that it was characteristic of the IVC. After the IVC’s decline, this population mixed with north- western groups with Steppe The formation of human populations ◥ ON OUR WEBSITE ancestry to form the “An- cestral North Indians”(ANI) in South and Central Asia Read the full article at http://dx.doi. and also mixed with south- org/10.1126/ eastern groups to form the Vagheesh M. Narasimhan et al. science.aat7487 “Ancestral South Indians” .................................................. (ASI), whose direct descend- RATIONALE: To elucidate the extent to which ically documented dispersal of domesticates ants today live in tribal groups in southern the major cultural transformations of farming, was accompanied by the spread of people from India. Mixtures of these two post-IVC groups— pastoralism, and shifts in the distribution of multiple centers of domestication. the ANI and ASI—drive the main gradient of languages in Eurasia were accompanied by The main population of the BMAC carried no genetic variation in South Asia today. movement of people, we report genome-wide ancestry from Steppe pastoralists and did not ancient DNA data from 523 individuals span- contribute substantially to later South Asians. CONCLUSION: Earlier work recorded mas- ning the last 8000 years, mostly from Central However, Steppe pastoralist ancestry appeared sive population movement from the Eurasian Downloaded from Asia and northernmost South Asia. in outlier individuals at BMAC sites by the turn Steppe into Europe early in the third millen- of the second millennium BCE around the same nium BCE, likely spreading Indo-European RESULTS: The movement of people follow- time as it appeared on the southern Steppe. languages. We reveal a parallel series of events ing the advent of farming resulted in genetic Using data from ancient individuals from the leading to the spread of Steppe ancestry to gradients across Eurasia that can be modeled Swat Valley of northernmost South Asia, we show South Asia, thereby documenting movements as mixtures of seven deeply divergent popula- that Steppe ancestry then integrated further south of people that were likely conduits for the http://science.sciencemag.org/ tions. A key gradient formed in southwestern in the first half of the second millennium BCE, spread of Indo-European languages.▪ Asia beginning in the Neolithic and continu- contributing up to 30% of the ancestry of modern ing into the Bronze Age, with more Anatolian groups in South Asia. The Steppe ancestry in farmer–related ancestry in the west and more South Asia has the same profile as that in Bronze The list of authors and affiliations is available in the full Iranian farmer–related ancestry in the east. Age Eastern Europe, tracking a movement of article online. Corresponding authors: Vagheesh M. Narasimhan (vagheesh@ This cline extended to the desert oases of people that affected both regions and that likely mail.harvard.edu); Nick Patterson ([email protected]); Central Asia and was the primary source of spread the unique features shared between Indo- Michael Frachetti ([email protected]); Ron Pinhasi (ron. ancestry in peoples of the Bronze Age Bactria Iranian and Balto-Slavic languages. [email protected]); David Reich ([email protected]. harvard.edu) Margiana Archaeological Complex (BMAC). The primary ancestral population of modern Cite this article as V. M. Narasimhan et al., Science 365, eaat7487 (2019). DOI: 10.1126/science.aat7487 This supports the idea that the archaeolog- South Asians is a mixture of people related to on September 5, 2019 The Impact of Yamnaya Steppe Pastoralists ~3000 BCE Yamnaya Other ancestry Location of the Flow of Yamnaya-derived ancestry initial formation of KAZAKHSTAN All arrows are approximate. Yamnaya ancestry is uncertain. Eurasian Steppe ~1700 BCE ~1700 BCE 0 800 km Yamnaya ASIA ~3300 BCE Ca sp EUROP E ia ~1700 BCE n E ~2700 BCE N IN S ~2400 BCE R L e E C a ~1000 BCE D N Sea ~2000 BCE O A ck E la M P B O Path by which R U Ancestral E this ancestry arrived North Indians ~2500 BCE in South Asia is 2000–1000 BCE M O uncertain. D E ATLANTIC R C N L I OCEAN IN N E D I A ea N ean S Mediterran Arabian Sea Ancestral ~2300 BCE South Indians AFRICA 2000–1000 BCE The Bronze Age spread of Yamnaya Steppe pastoralist ancestry into two subcontinents—Europe and South Asia. Pie charts reflect the proportion of Yamnaya ancestry, and dates reflect the earliest available ancient DNA with Yamnaya ancestry in each region. Ancient DNA has not yet been found for the ANI and ASI, so for these the range is inferred statistically. Narasimhan et al., Science 365, 999 (2019) 6 September 2019 1of1 RESEARCH ◥ the Iranian plateau to the desert oases south of RESEARCH ARTICLE the Eurasian Steppe was accompanied by move- ments of people or adoption of ideas from neigh- boring groups (2–4). For the urban communities HUMAN EVOLUTION of the Bactria Margiana Archaeological Complex (BMAC) in the Bronze Age, we assessed whether the people buried in its cemeteries descended The formation of human populations directly from earlier smaller-scale food producers, andwealsodocumentedtheirgenetichetero- in South and Central Asia geneity (5). Farther to the north and east, we showed that the Early Bronze Age spread of crops and domesticated animals between Southwest 1 † 2,3 † 4,5‡ 1,2‡ Vagheesh M. Narasimhan * , Nick Patterson * , Priya Moorjani , Nadin Rohland , Asia and eastern Eurasia along the Inner Asian 1 1,2,6‡ 1 1,7 Rebecca Bernardos , Swapan Mallick , Iosif Lazaridis , Nathan Nakatsuka , Mountain Corridor (6) was accompanied by 1 1 1,8 9 10 Iñigo Olalde , Mark Lipson , Alexander M. Kim , Luca M. Olivieri , Alfredo Coppa , movements of people. Finally, we examined 9,11 12 13 14,15,16 Massimo Vidale , James Mallory , Vyacheslav Moiseyev , Egor Kitov , when descendants of the Yamnaya, who spread 17 1,6 18 1,6 Janet Monge , Nicole Adamski , Neel Alex , Nasreen Broomandkhoshbacht §, across the Eurasian Steppe beginning around 19,20 1,6 19,21,22 Francesca Candilio , Kimberly Callan , Olivia Cheronet , 3300 BCE (7–9), began to appear in Central Asia Brendan J. Culleton23, Matthew Ferry1,6, Daniel Fernandes19,21,22,24, Suzanne Freilich22, south of the Steppe. Beatriz Gamarra19,21,25||¶, Daniel Gaudio19,21, Mateja Hajdinjak26, Éadaoin Harney1,6,27, In northernmost South Asia, we report a time Thomas K. Harper28, Denise Keating19, Ann Marie Lawson1,6, Matthew Mah1,2,6, transect of >100 individuals beginning ~1200 BCE, Downloaded from Kirsten Mandl22, Megan Michel1,6#**, Mario Novak19,29, Jonas Oppenheimer1,6††, which we coanalyze along with modern data from Niraj Rai30,31, Kendra Sirak1,19,32, Viviane Slon26, Kristin Stewardson1,6, hundreds of present-day South Asian groups, as Fatma Zalzala1,6, Zhao Zhang1, Gaziz Akhatov15, Anatoly N. Bagashev33, well as ancient DNA from neighboring regions Alessandra Bagnera9, Bauryzhan Baitanayev15, Julio Bendezu-Sarmiento34, (10). Previous analyses place the majority of Arman A. Bissembaev15,35, Gian Luca Bonora36, Temirlan T. Chargynov37, present-day South Asians along a genetic cline (11) 38 39 38 40 Tatiana Chikisheva , Petr K. Dashkovskiy , Anatoly Derevianko , Miroslav Dobeš , that can be modeled as having arisen from mixture http://science.sciencemag.org/ Katerina Douka41,42, Nadezhda Dubova14, Meiram N. Duisengali35, Dmitry Enshin33, of two highly divergent populations after around Andrey Epimakhov43,44, Alexey V. Fribus45, Dorian Fuller46,47, Alexander Goryachev33, 4000 years ago: the Ancestral North Indians (ANI), who harbor large proportions of ancestry Andrey Gromov13, Sergey P. Grushin48, Bryan Hanks49, Margaret Judd49, related to West Eurasians, and the Ancestral South Erlan Kazizov15, Aleksander Khokhlov50, Aleksander P. Krygin51, Elena Kupriyanova52, Indians (ASI), who are much less closely related to Pavel Kuznetsov50, Donata Luiselli53, Farhod Maksudov54, Aslan M. Mamedov55, West Eurasians (12). We leveraged ancient DNA to Talgat B. Mamirov15, Christopher Meiklejohn56, Deborah C. Merrett57, 9,58 50 54,59 41 place constraints on the genetic structure of the Roberto Micheli , Oleg Mochalov , Samariddin Mustafokulov , Ayushi Nayak , ANI and ASI and, in conjunction with other lines 60 61 33 62 Davide Pettener , Richard Potts , Dmitry Razhev , Marina Rykun , of evidence, to make inferences about when and 60 63 64 Stefania Sarno , Tatyana M. Savenkova , Kulyan Sikhymbaeva , where they formed. By modeling modern South 33 37 38 Sergey M. Slepchenko , Oroz A. Soltobaev , Nadezhda Stepanova , Asians along with ancient individuals from sites on September 5, 2019 13,65 66 22,67 Svetlana Svyatko , Kubatbek Tabaldiev , Maria Teschler-Nicola , in cultural contact with the Indus Valley Civili- Alexey A. Tishkin68, Vitaly V. Tkachev69, Sergey Vasilyev14,70, Petr Velemínský71, zation (IVC), we inferred a likely genetic signa- Dmitriy Voyakin15,72, Antonina Yermolayeva15, Muhammad Zahir41,73, ture for people of the IVC that reached its Valery S. Zubkov74, Alisa Zubova13, Vasant S. Shinde75, Carles Lalueza-Fox76, maturity in northwestern South Asia between Matthias Meyer26, David Anthony77, Nicole Boivin41‡, Kumarasamy Thangaraj30‡, 2600and1900BCE.WealsoexaminedwhenSteppe Douglas J. Kennett23,28,78‡, Michael Frachetti79,80†, pastoralist–derived ancestry (9) mixed into groups Ron Pinhasi19,22†, David Reich1,2,6,81† in South Asia, and placed constraints on whether Steppe-related ancestry or Iranian-related ances- By sequencing 523 ancient humans, we show that the primary source of ancestry in try is more plausibly associated with the spread modern South Asians is a prehistoric genetic gradient between people related to early of Indo-European languages in South Asia. hunter-gatherers of Iran and Southeast Asia. After the Indus Valley Civilization’s Dataset and analysis strategy decline, its people mixed with individuals in the southeast to form one of the two main ancestral populations of South Asia, whose direct descendants live in southern India.
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