The Genetic Origin of the Indo-Europeans
Authors: Lazaridis Iosif , Patterson Nick , Anthony David , Vyazov Leonid , Fournier Romain , Ringbauer Harald , Olalde Iñigo , Khokhlov Alexander A. , Kitov Egor P. , Shishlina Natalia I. , Ailincăi Sorin C. , Agapov Danila S. , Agapov Sergey A. , Batieva Elena , Bauyrzhan Baitanayev , Bereczki Zsolt , Buzhilova Alexandra , Changmai Piya , Chizhevsky Andrey A. , Ciobanu Ion , Constantinescu Mihai , Csányi Marietta , Dani János , Dashkovskiy Peter K. , Évinger Sándor , Faifert Anatoly , Flegontov Pavel N. , Frînculeasa Alin , Frînculeasa Mădălina N. , Hajdu Tamás , Higham Tom , Jarosz Paweł , Jelínek Pavol , Khartanovich Valeri I. , Kirginekov Eduard N. , Kiss Viktória , Kitova Alexandera , Kiyashko Alexeiy V. , Koledin Jovan , Korolev Arkady , Kosintsev Pavel , Kulcsár Gabriella , Kuznetsov Pavel , Magomedov Rabadan , Malikovich Mamedov Aslan , Melis Eszter , Moiseyev Vyacheslav , Molnár Erika , Monge Janet , Negrea Octav , Nikolaeva Nadezhda A. , Novak Mario , Ochir-Goryaeva Maria , Pálfi György , Popovici Sergiu , Rykun Marina P. , Savenkova Tatyana M. , Semibratov Vladimir P. , Seregin Nikolai N. , Šefčáková Alena , Serikovna Mussayeva Raikhan , Shingiray Irina , Shirokov Vladimir N. , Simalcsik Angela , Sirak Kendra , Solodovnikov Konstantin N. , Tárnoki Judit , Tishkin Alexey A. , Trifonov Viktov , Vasilyev Sergey , Akbari Ali , Brielle Esther S. , Callan Kim , Candilio Francesca , Cheronet Olivia , Curtis Elizabeth , Flegontova Olga , Iliev Lora , Kearns Aisling , Keating Denise , Lawson Ann Marie , Mah Matthew , Micco Adam , Michel Megan , Oppenheimer Jonas , Qiu Lijun , Workman J. Noah , Zalzala Fatma , Szécsényi-Nagy Anna , Palamara Pier Francesco , Mallick Swapan , Rohland Nadin , Pinhasi Ron , Reich David 
Published: April 2024

The Yamnaya archaeological complex appeared around 3300BCE across the steppes north of the Black and Caspian Seas, and by 3000BCE reached its maximal extent from Hungary in the west to Kazakhstan in the east. To localize the ancestral and geographical origins of the Yamnaya among the diverse Eneolithic people that preceded them, we studied ancient DNA data from 428 individuals of which 299 are reported for the first time, demonstrating three previously unknown Eneolithic genetic clines. First, a “Caucasus-Lower Volga” (CLV) Cline suffused with Caucasus hunter-gatherer (CHG) ancestry extended between a Caucasus Neolithic southern end in Neolithic Armenia, and a steppe northern end in Berezhnovka in the Lower Volga. Bidirectional gene flow across the CLV cline created admixed intermediate populations in both the north Caucasus, such as the Maikop people, and on the steppe, such as those at the site of Remontnoye north of the Manych depression. CLV people also helped form two major riverine clines by admixing with distinct groups of European hunter-gatherers. A “Volga Cline” was formed as Lower Volga people mixed with upriver populations that had more Eastern hunter-gatherer (EHG) ancestry, creating genetically hyper-variable populations as at Khvalynsk in the Middle Volga. A “Dnipro Cline” was formed as CLV people bearing both Caucasus Neolithic and Lower Volga ancestry moved west and acquired Ukraine Neolithic hunter-gatherer (UNHG) ancestry to establish the population of the Serednii Stih culture from which the direct ancestors of the Yamnaya themselves were formed around 4000BCE. This population grew rapidly after 3750-3350BCE, precipitating the expansion of people of the Yamnaya culture who totally displaced previous groups on the Volga and further east, while admixing with more sedentary groups in the west. CLV cline people with Lower Volga ancestry contributed four fifths of the ancestry of the Yamnaya, but also, entering Anatolia from the east, contributed at least a tenth of the ancestry of Bronze Age Central Anatolians, where the Hittite language, related to the Indo-European languages spread by the Yamnaya, was spoken. We thus propose that the final unity of the speakers of the “Proto-Indo-Anatolian” ancestral language of both Anatolian and Indo-European languages can be traced to CLV cline people sometime between 4400-4000 BCE.Abstract FigureSummary Figure:The origin of Indo-Anatolian and Indo-European languages.Genetic reconstruction of the ancestry of Pontic-Caspian steppe and West Asian populations points to the North Caucasus-Lower Volga area as the homeland of Indo-Anatolian languages and to the Serednii Stih archaeological culture of the Dnipro-Don area as the homeland of Indo-European languages. The Caucasus-Lower Volga people had diverse distal roots, estimated using theqpAdmsoftware on the left barplot, as Caucasus hunter-gatherer (purple), Central Asian (red), Eastern hunter-gatherer (pink), and West Asian Neolithic (green). Caucasus-Lower Volga expansions, estimated usingqpAdmon the right barplot as disseminated Caucasus Neolithic (blue)-Lower Volga Eneolithic (orange) proximal ancestries, mixing with the inhabitants of the North Pontic region (yellow), Volga region (yellow), and West Asia (green).