The genetic origin of the Indo-Europeans
Iosif Lazaridis, Nick Patterson, David W. Anthony, Leonid Vyazov, Romain Fournier, Harald Ringbauer, Íñigo Olalde, Alexander A. Khokhlov, Еgor Kitov, Natalia Shishlina, Sorin Ailincăi, Danila S. Agapov, Sergey A. Agapov, Elena Batieva, Baitanayev Bauyrzhan, Zsolt Bereczki, Alexandra Buzhilova, Piya Changmai, Andrey A. Chizhevsky, Ion Ciobanu, Mihai Constantinescu, Marietta Csányi, János Dani, Петр Константинович Дашковский, Sándor Évinger, Anatoly Faifert, Pavel Flegontov, Alin Frînculeasa, Mădălina N. Frînculeasa, Tamás Hajdu, Thomas Higham, Paweł Jarosz, Pavol Jelínek, Valeri Khartanovich, Eduard N. Kirginekov, Viktória Kiss, Alexandera Kitova, Alexeiy V. Kiyashko, Jovan Koledin, Arkady Ivanovich Korolev, П. А. Косинцев, Gabriella Kulcsár, Pavel Kuznetsov, R. A. Magomedov, Aslan M. Mamedov, Eszter Melis, Vyacheslav Moiseyev, Erika Molnár, Janet Monge, Octav Negrea, Н. А. Николаева, Mario Novak, Maria A. Ochir-Goryaeva, György Pálfi, Sergiu Popovici, Marina P. Rykun, Tatyana M. Savenkova, Vladimir P. Semibratov, N. Seregin, Alena Šefčáková, Raikhan S. Mussayeva, Irina Shingiray, Vladimir N. Shirokov, Angela Simalcsik, Kendra Sirak, K.N. Solodovnikov, Judit Tárnoki, А. Тишкин, Viktor Trifonov, Sergey Vasilyev, Ali Akbari, Esther S. Brielle, Kimberly Callan, Francesca Candilio, Olivia Cheronet, Elizabeth Curtis, Olga Flegontova, Lora Iliev, Aisling Kearns, Denise Keating, Ann Marie Lawson, Matthew Mah, Adam Micco, Megan Michel, Jonas Oppenheimer, Lijun Qiu, J. Noah Workman, Fatma Zalzala, Anna Szécsényi‐Nagy, Pier Francesco Palamara, Swapan Mallick, Nadin Rohland, Ron Pinhasi, David Reich
Abstract
The Yamnaya archaeological complex appeared around 3300 bc across the steppes north of the Black and Caspian Seas, and by 3000 bc it reached its maximal extent, ranging from Hungary in the west to Kazakhstan in the east. To localize Yamnaya origins among the preceding Eneolithic people, we assembled ancient DNA from 435 individuals, demonstrating three genetic clines. A Caucasus–lower Volga (CLV) cline suffused with Caucasus hunter-gatherer1 ancestry extended between a Caucasus Neolithic southern end and a northern end at Berezhnovka along the lower Volga river. Bidirectional gene flow created intermediate populations, such as the north Caucasus Maikop people, and those at Remontnoye on the steppe. The Volga cline was formed as CLV people mixed with upriver populations of Eastern hunter-gatherer2 ancestry, creating hypervariable groups, including one at Khvalynsk. The Dnipro cline was formed when CLV people moved west, mixing with people with Ukraine Neolithic hunter-gatherer ancestry3 along the Dnipro and Don rivers to establish Serednii Stih groups, from whom Yamnaya ancestors formed around 4000 bc and grew rapidly after 3750–3350 bc. The CLV people contributed around four-fifths of the ancestry of the Yamnaya and, entering Anatolia, probably from the east, at least one-tenth of the ancestry of Bronze Age central Anatolians, who spoke Hittite4,5. We therefore propose that the final unity of the speakers of ‘proto-Indo-Anatolian’, the language ancestral to both Anatolian and Indo-European people, occurred in CLV people some time between 4400 bc and 4000 bc. Ancient DNA reveals how the explosive expansion of Yamnaya steppe pastoralists began with a small community north of the Black Sea speaking ancestral Indo-European, and detects genetic links with Anatolian speakers, stemming from a common Indo-Anatolian homeland in the North Caucasus–lower Volga region.