Yamnaya culture explained

Yamnaya culture
Map:Yamnaya culture.jpg
Region:Pontic–Caspian steppe in Europe
Period:Copper Age, Bronze Age
Dates:2600 BCE
Precededby:Sredny Stog culture, Samara culture, Khvalynsk culture, Dnieper–Donets culture, Repin culture, Maykop culture, Cucuteni–Trypillia culture, Cernavodă culture, Usatove culture, Novosvobodnaya culture
Followedby:
    West: Catacomb culture, Vučedol cultureEast: Poltavka culture
    Definedby:Vasily Gorodtsov

    The Yamnaya culture or the Yamna culture, also known as the Pit Grave culture or Ochre Grave culture, is a late Copper Age to early Bronze Age archaeological culture of the region between the Southern Bug, Dniester, and Ural rivers (the Pontic–Caspian steppe), dating to 3300–2600 BCE. It was discovered by Vasily Gorodtsov following his archaeological excavations near the Donets River in 1901–1903. Its name derives from its characteristic burial tradition: Russian: Я́мная (romanization:) is a Russian adjective that means 'related to pits ', as these people used to bury their dead in tumuli (kurgans) containing simple pit chambers. Research in recent years has found that Mikhaylovka, in lower Dnieper river, Ukraine, formed the Core Yamnaya culture (c. 3600–3400 BCE).[1]

    The Yamnaya economy was based upon animal husbandry, fishing, and foraging, and the manufacture of ceramics, tools, and weapons. The people of the Yamnaya culture lived primarily as nomads, with a chiefdom system and wheeled carts and wagons that allowed them to manage large herds. They are also closely connected to Final Neolithic cultures, which later spread throughout Europe and Central Asia, especially the Corded Ware people and the Bell Beaker culture, as well as the peoples of the Sintashta, Andronovo, and Srubnaya cultures. Back migration from Corded Ware also contributed to Sintashta and Andronovo. In these groups, several aspects of the Yamnaya culture are present.[2] Yamnaya material culture was very similar to the Afanasevo culture of South Siberia, and the populations of the two cultures are genetically indistinguishable. This suggests that the Afanasevo culture may have originated from the migration of Yamnaya groups to the Altai region or, alternatively, that both cultures developed from an earlier shared cultural source.[3]

    Genetic studies have suggested that the people of the Yamnaya culture can be modelled as a genetic admixture between a population related to Eastern European Hunter-Gatherers (EHG)[4] and people related to hunter-gatherers from the Caucasus (CHG) in roughly equal proportions,[5] an ancestral component which is often named "Steppe ancestry", with additional admixture from Anatolian, Levantine, or Early European farmers.[6] Genetic studies also indicate that populations associated with the Corded Ware, Bell Beaker, Sintashta, and Andronovo cultures derived large parts of their ancestry from the Yamnaya or a closely related population.[7]

    According to the widely-accepted Kurgan hypothesis of Marija Gimbutas, the people that produced the Yamnaya culture spoke a stage of the Proto Indo-European language, which later spread eastwards and westwards as part of the Indo-European migrations.

    Origins

    See also: Kurgan hypothesis and Marija Gimbutas. The Yamnaya culture was defined by Vasily Gorodtsov in order to differentiate it from the Catacomb and Srubnaya cultures that existed in the area, but were considered to be of a later period. Due to the time interval to the Yamnaya culture, and the reliance on archaeological findings, debate as to its origin is ongoing. In 1996, Pavel Dolukhanov suggested that the emergence of the Pit-Grave culture represents a social development of various different local Bronze Age cultures, thus representing "an expression of social stratification and the emergence of chiefdom-type nomadic social structures" which in turn intensified inter-group contacts between essentially heterogeneous social groups.

    The origin of the Yamnaya culture continues to be debated, with proposals for its origins pointing to both the Khvalynsk and Sredny Stog cultures. The Khvalynsk culture (4700–3800 BCE) (middle Volga) and the Don-based Repin culture (–3300 BCE) in the eastern Pontic-Caspian steppe, and the closely related Sredny Stog culture (–3500 BCE) in the western Pontic-Caspian steppe, preceded the Yamnaya culture (3300–2500 BCE).

    Further efforts to pinpoint the location came from Anthony (2007), who suggested that the Yamnaya culture (3300–2600 BCE) originated in the DonVolga area at, preceded by the middle Volga-based Khvalynsk culture and the Don-based Repin culture (–3300 BCE), arguing that late pottery from these two cultures can barely be distinguished from early Yamnaya pottery. Earlier continuity from eneolithic but largely hunter-gatherer Samara culture and influences from the more agricultural Dnieper–Donets II are apparent.

    He argues that the early Yamnaya horizon spread quickly across the Pontic–Caspian steppes between and 3200 BCE:

    Alternatively, Parpola (2015) relates both the Corded ware culture and the Yamnaya culture to the late Trypillia (Tripolye) culture. He hypothesizes that "the Tripolye culture was taken over by PIE speakers by c. 4000 BCE," and that in its final phase the Trypillian culture expanded to the steppes, morphing into various regional cultures which fused with the late Sredny Stog (Serednii Stih) pastoralist cultures, which, he suggests, gave rise to the Yamnaya culture. Dmytro Telegin viewed Sredny Stog and Yamna as one cultural continuum and considered Sredny Stog to be the genetic foundation of the Yamna.[8]

    The Yamnaya culture was succeeded in its western range by the Catacomb culture (2800–2200 BCE); in the east, by the Poltavka culture (2700–2100 BCE) at the middle Volga. These two cultures were followed by the Srubnaya culture (18th–12th century BCE).

    Characteristics

    The Yamnaya culture was nomadic or semi-nomadic, with some agriculture practiced near rivers, and a few fortified sites, the largest of which is Mikhaylivka.

    Characteristic of the culture are the burials in pit graves surmounted by kurgans (tumuli), often accompanied by animal offerings. Some graves contain large anthropomorphic stelae, with carved human heads, arms, hands, belts, and weapons. The bodies were placed in a supine position with bent knees and covered in ochre. Some kurgans contained "stratified sequences of graves". Kurgan burials may have been rare, and were perhaps reserved for special adults, who were predominantly, but not necessarily, male.[9] Status and gender are marked by grave goods and position, and in some areas, elite individuals are buried with complete wooden wagons.[10] Grave goods are more common in eastern Yamnaya burials, which are also characterized by a higher proportion of male burials and more male-centred rituals than western areas.

    The Yamnaya culture had and used two-wheeled carts and four-wheeled wagons, which are thought to have been oxen-drawn at this time, and there is evidence that they rode horses.[11] [12] For instance, several Yamnaya skeletons exhibit specific characteristics in their bone morphology that may have been caused by long-term horseriding.

    Metallurgists and other craftsmen are given a special status in Yamnaya society, and metal objects are sometimes found in large quantities in elite graves. New metalworking technologies and weapon designs are used.

    Stable isotope ratios of Yamna individuals from the Dnipro Valley suggest the Yamnaya diet was terrestrial protein based with insignificant contribution from freshwater or aquatic resources.[13] Anthony speculates that the Yamnaya ate meat, milk, yogurt, cheese, and soups made from seeds and wild vegetables, and probably consumed mead.

    Mallory and Adams suggest that Yamnaya society may have had a tripartite structure of three differentiated social classes, although the evidence available does not demonstrate the existence of specific classes such as priests, warriors, and farmers.[14]

    Archaeogenetics

    According to Jones et al. (2015) and Haak et al. (2015), autosomal tests indicate that the Yamnaya people were the result of a genetic admixture between two different hunter-gatherer populations: distinctive "Eastern Hunter-Gatherers" (EHG), from Eastern Europe, with high affinity to the Mal'ta–Buret' culture or other, closely related people from Siberia and a population of "Caucasus hunter-gatherers" (CHG) who probably arrived from the Caucasus or Iran. Each of those two populations contributed about half the Yamnaya DNA. This admixture is referred to in archaeogenetics as Western Steppe Herder (WSH) ancestry.

    Admixture between EHGs and CHGs is believed to have occurred on the eastern Pontic-Caspian steppe starting around 5,000 BCE, while admixture with Early European Farmers (EEF) happened in the southern parts of the Pontic-Caspian steppe sometime later. More recent genetic studies have found that the Yamnaya were a mixture of EHGs, CHGs, and to a lesser degree Anatolian farmers and Levantine farmers, but not EEFs from Europe due to lack of WHG DNA in the Yamnaya. This occurred in two distinct admixture events from West Asia into the Pontic-Caspian steppe.[15] [16]

    Haplogroup R1b, specifically the Z2103 subclade of R1b-L23, is the most common Y-DNA haplogroup found among the Yamnaya specimens. This haplogroup is rare in Western Europe and mainly exists in Southeastern Europe today. Additionally, a minority are found to belong to haplogroup I2. They are found to belong to a wider variety of West Eurasian mtDNA haplogroups, including U, T, and haplogroups associated with Caucasus Hunter-Gatherers and Early European Farmers.[17] A small but significant number of Yamnaya kurgan specimens from Northern Ukraine carried the East Asian mtDNA haplogroup C4.[18] [19]

    People of the Yamnaya culture are believed to have had mostly brown eye colour, light to intermediate skin, and brown hair colour, with some variation.[20] [21]

    Some Yamnaya individuals are believed to have carried a mutation to the KITLG gene associated with blond hair, as several individuals with Steppe ancestry are later found to carry this mutation. The Ancient North Eurasian Afontova Gora group, who contributed significant ancestry to Western Steppe Herders, are believed to be the source of this mutation.[22] A study in 2015 found that Yamnaya had the highest ever calculated genetic selection for height of any of the ancient populations tested.[23] [24] It has been hypothesized that an allele associated with lactase persistence (conferring lactose tolerance into adulthood) was brought to Europe from the steppe by Yamnaya-related migrations.[25] [26] [27] [28]

    A 2022 study by Lazaridis et al. found that the typical phenotype among the Yamnaya population was brown eyes, brown hair, and intermediate skin colour. None of their Yamnaya samples were predicted to have either blue eyes or blond hair, in contrast with later Steppe groups in Russia and Central Asia, as well as the Bell Beaker culture in Europe, who did carry these phenotypes in high proportions.

    The geneticist David Reich has argued that the genetic data supports the likelihood that the people of the Yamnaya culture were a "single, genetically coherent group" who were responsible for spreading many Indo-European languages.[29] Reich's group recently suggested that the source of Anatolian and Indo-European subfamilies of the Proto-Indo-European (PIE) language may have been in west Asia and the Yamna were responsible for the dissemination of the latter.[30] Reich also argues that the genetic evidence shows that Yamnaya society was an oligarchy dominated by a small number of elite males.[31]

    The genetic evidence for the extent of the role of the Yamnaya culture in the spread of Indo-European languages has been questioned by Russian archaeologist Leo Klejn[32] and Balanovsky et al.,[33] who note a lack of male haplogroup continuity between the people of the Yamnaya culture and the contemporary populations of Europe. Klejn has also suggested that the autosomal evidence does not support a Yamnaya migration, arguing that Western Steppe Herder ancestry in both contemporary and Bronze Age samples is lowest around the Danube in Hungary, near the western limits of the Yamnaya culture, and highest in Northern Europe, which Klejn argues is the opposite of what would be expected if the geneticists' hypothesis is correct.

    Language

    Marija Gimbutas identified the Yamnaya culture with the late Proto-Indo-Europeans (PIE) in her Kurgan hypothesis. In the view of David Anthony, the Pontic-Caspian steppe is the strongest candidate for the Urheimat (original homeland) of the Proto-Indo-European language, citing evidence from linguistics and genetics[34] which suggests that the Yamnaya culture may be the homeland of the Indo-European languages, with the possible exception of the Anatolian languages. On the other hand, Colin Renfrew has argued for a Near Eastern origin of the earliest Indo-European speakers.[35]

    According to David W. Anthony, the genetic evidence suggests that the leading clans of the Yamnaya were of EHG (Eastern European hunter-gatherer) and WHG (Western European hunter-gatherer) paternal origin and implies that the Indo-European languages were the result of "a dominant language spoken by EHGs that absorbed Caucasus-like elements in phonology, morphology, and lexicon." It has also been suggested that the PIE language evolved through trade interactions in the circum-Pontic area in the 4th millennium BCE, mediated by the Yamna predecessors in the North Pontic steppe.[36]

    Guus Kroonen et al. 2022 found that the "basal Indo-European stage", also known as Indo-Anatolian or Pre-Proto-Indo-European language, largely but not totally, lacked agricultural-related vocabulary, and only the later "core Indo-European languages" saw an increase in agriculture-associated words. According to them, this fits a homeland of early core Indo-European within the westernmost Yamnaya horizon, around and west of the Dnieper, while its basal stage, Indo-Anatolian, may have originated in the Sredny Stog culture, as opposed to the eastern Yamnaya horizon. The Corded Ware culture may have acted as major source for the spread of later Indo-European languages, including Indo-Iranian, while Tocharian languages may have been mediated via the Catacomb culture. They also argue that this new data contradicts a possible earlier origin of Pre-Proto-Indo-European among agricultural societies South of the Caucasus, rather "this may support a scenario of linguistic continuity of local non-mobile herders in the Lower Dnieper region and their genetic persistence after their integration into the successive and expansive Yamnaya horizon". Furthermore the authors mention that this scenario can explain the difference in paternal haplogroup frequency between the Yamnaya and Corded Ware cultures, while both sharing similar autosomal DNA ancestry.[37]

    Yamnaya-related migrations

    See also: Indo-European migrations.

    Western Europe

    Genetic studies have found that Yamnaya autosomal characteristics are very close to the Corded Ware culture people, with up to 75% Yamnaya-like ancestry in the DNA of Corded Ware skeletons from Central and Eastern Europe.[38] Yamnaya–related ancestry is found in the DNA of modern Central, and Northern Europeans (c. 38.8–50.4 %), and is also found in lower levels in present-day Southern Europeans (c. 18.5–32.6 %), Sardinians (c. 2.4–7.1 %), and Sicilians (c. 5.9–11.6 %).

    However, according to Heyd, et al. (2023), the specific paternal DNA haplogroup that is most commonly found in male Yamnaya specimens cannot be found in modern Western Europeans, or in males from the nearby Corded Ware culture. This makes it unlikely that the Corded Ware culture can be directly descended from the Yamnaya culture, at least along the paternal line.[39]

    Autosomal tests also indicate that the Yamnaya are the vector for "Ancient North Eurasian" admixture into Europe. "Ancient North Eurasian" is the name given in literature to a genetic component that represents descent from the people of the Mal'ta–Buret' culture or a population closely related to them. That genetic component is visible in tests of the Yamnaya people as well as modern-day Europeans.

    Eastern Europe and Finland

    In the Baltic, Jones et al. (2017) found that the Neolithic transition – the passage from a hunter-gatherer economy to a farming-based economy – coincided with the arrival en masse of individuals with Yamnaya-like ancestry. This is different from what happened in Western and Southern Europe, where the Neolithic transition was caused by a population that came from Anatolia, with Pontic steppe ancestry being detected from only the late Neolithic onward.

    Per Haak et al. (2015), the Yamnaya contribution in the modern populations of Eastern Europe ranges from 46.8% among Russians to 42.8% in Ukrainians. Finland has the highest Yamnaya contributions in all of Europe (50.4%).[40]

    Central and South Asia

    See also: Sintashta culture.

    Studies also point to the strong presence of Yamnaya descent in the current nations of South Asia, especially in groups that are referred to as Indo-Aryans. According to Pathak et al. (2018), the "North-Western Indian & Pakistani" populations (PNWI) showed significant Middle-Late Bronze Age Steppe (Steppe_MLBA) ancestry along with Yamnaya Early-Middle Bronze Age (Steppe_EMBA) ancestry, but the Indo-Europeans of Gangetic Plains and Dravidian people only showed significant Yamnaya (Steppe_EMBA) ancestry and no Steppe_MLBA. The study also noted that ancient south Asian samples had significantly higher Steppe_MLBA than Steppe_EMBA (or Yamnaya). The study identified the Rors and Jats as the population in South Asia with the highest proportion of Steppe ancestry. Lazaridis et al. (2016) estimated (6.5–50.2 %) steppe-related admixture in South Asians, though the proportion of Steppe ancestry varies widely across ethnic groups. According to Narasimhan et al. (2019), the Yamnaya-related ancestry, termed Western_Steppe_EMBA, that reached central and south Asia was not the initial expansion from the steppe to the east, but a secondary expansion that involved a group possessing ~67% Western_Steppe_EMBA ancestry and ~33% ancestry from the European cline. This group included people similar to that of Corded Ware, Srubnaya, Petrovka, and Sintashta. Moving further east in the central steppe, it acquired ~9% ancestry from a group of people that possessed West Siberian Hunter Gatherer ancestry, thus forming the Central Steppe MLBA cluster, which is the primary source of steppe ancestry in South Asia, contributing up to 30% of the ancestry of the modern groups in the region.

    According to Unterländer et al. (2017), all Iron Age Scythian Steppe nomads can best be described as a mixture of Yamnaya-related ancestry and an East Asian-related component, which most closely corresponds to the modern North Siberian Nganasan people of the lower Yenisey River, to varying degrees, but generally higher among Eastern Scythians.

    See also

    References

    Sources

    External links

    Notes and References

    1. Reich, David, (24 April, 2024). "The Genetic Origin of the Indo-Europeans", in: The Transfomation of Europe in the Third Millennium BC, International Conference, HUN-REN RCH Institute of Archaeology, Budapest.
    2. Yamnayan cultural aspects, for example, were horse-riding, burial styles, and to some extent the pastoralist economy.
    3. Hermes . Taylor R. . Tishkin . Alexey A. . Kosintsev . Pavel A. . Stepanova . Nadezhda F. . Krause-Kyora . Ben . Makarewicz . Cheryl A. . 1 December 2020 . Mitochondrial DNA of domesticated sheep confirms pastoralist component of Afanasievo subsistence economy in the Altai Mountains (3300-2900 cal BC). Archaeological Research in Asia . en . 24 . 100232 . 10.1016/j.ara.2020.100232 . 225136827 . 2352-2267.
    4. The Eastern European hunter-gatherers were themselves mostly descended from ancient North Eurasians, related to the palaeolithic Mal'ta–Buret' culture.
    5. News: Europe's fourth ancestral 'tribe' uncovered. BBC . 16 November 2015 .
    6. Wang. Chuan-Chao. Reinhold. Sabine. Kalmykov. Alexey. Wissgott. Antje. Brandt. Guido. Jeong. Choongwon. Cheronet. Olivia. Ferry. Matthew. Harney. Eadaoin. Keating. Denise. Mallick. Swapan. 4 February 2019. Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions. Nature Communications. 10. 1. 590. 10.1038/s41467-018-08220-8. 30713341. 2041-1723. 6360191. 2019NatCo..10..590W.
    7. News: Ann . Gibbons . 10 June 2015 . Nomadic herders left a strong genetic mark on Europeans and Asians . AAAS . Science.
    8. Book: Telegin, D. Y. . Serednʹo-stohivsʹka kulʹtura epokhy midi . Naukova Dumka . 1973 . Kyiv . 147.
    9. Book: Anthony, David . The Horse, the Wheel, and Language. 1102387902. 328–329.
    10. Book: J., Harrison, Richard . The transformation of Europe in the third millennium BC: the example of "Le Petit-Chasseur I + II" (Sion, Valais, Switzerland). 196. 718304072.
    11. Science Advances . 9 . 9 . 2023 . First bioanthropological evidence for Yamnaya horsemanship . Trautmann . Martin . eade2451 . 10.1126/sciadv.ade2451. 36867690 . 10954216 . 2023SciA....9E2451T . 257334477 . 10138/356326 . free .
    12. Book: P., Mallory, J.. 2003 . 1989. Thames and Hudson. 0-500-27616-1. 213. 886668216.
    13. Book: Gerling, Claudia . Prehistoric Mobility and Diet in the West Eurasian Steppes 3500 to 300 BC: An Isotopic Approach . 1 July 2015 . De Gruyter . 978-3-11-031121-1 . en . 10.1515/9783110311211.
    14. Book: Encyclopedia of Indo-European culture. 1997. Fitzroy Dearborn. J. P. Mallory . Douglas Q. Adams. 1-884964-98-2. London. 653. 37931209.
    15. Lazaridis . Iosif . Alpaslan-Roodenberg . Songül . Acar . Ayşe . Açıkkol . Ayşen . Agelarakis . Anagnostis . Aghikyan . Levon . Akyüz . Uğur . Andreeva . Desislava . Andrijašević . Gojko . Antonović . Dragana . Armit . Ian . Atmaca . Alper . Avetisyan . Pavel . Aytek . Ahmet İhsan . Bacvarov . Krum . 26 August 2022 . The genetic history of the Southern Arc: A bridge between West Asia and Europe . Science . 377 . 6609 . eabm4247 . 10.1126/science.abm4247 . 36007055 . 10064553 . 251843620 . 0036-8075. free .
    16. Chintalapati . Manjusha . Patterson . Nick . Moorjani . Priya . 30 May 2022 . Perry . George H . The spatiotemporal patterns of major human admixture events during the European Holocene . eLife . 11 . e77625 . 10.7554/eLife.77625 . 35635751 . 9293011 . 2050-084X . free .
    17. Book: Eske, Allentoft . Morten E. Pokutta . Dalia Willerslev . Population genomics of Bronze Age Eurasia. 2015. University of Copenhagen, Denmark. 1234973657.
    18. Pilipenko . Aleksandr S. . Trapezov . Rostislav O. . Cherdantsev . Stepan V. . Babenko . Vladimir N. . Nesterova . Marina S. . Pozdnyakov . Dmitri V. . Molodin . Vyacheslav I. . Polosmak . Natalia V. . Maternal genetic features of the Iron Age Tagar population from Southern Siberia (1st millennium BC) . PLOS ONE . Public Library of Science (PLoS) . 13 . 9 . 20 September 2018 . 1932-6203 . 10.1371/journal.pone.0204062 . e0204062. free . 30235269 . 6147448 . 2018PLoSO..1304062P .
    19. Nikitin . Alexey G . Ivanova . Svetlana . Kiosak . Dmytro . Badgerow . Jessica . Pashnick . Jeff . Subdivisions of haplogroups U and C encompass mitochondrial DNA lineages of Eneolithic–Early Bronze Age Kurgan populations of western North Pontic steppe . Journal of Human Genetics . Springer Science and Business Media LLC . 62 . 6 . 2 February 2017 . 1434-5161 . 10.1038/jhg.2017.12 . 605–613. 28148921 . 7459815 .  "In the 12 successfully haplotyped specimens, 75% of mtDNA lineages consisted of west Eurasian haplogroup U and its U4 and U5 sublineages. Furthermore, we identified a subgroup of east Eurasian haplogroup C in two representatives of the Yamna culture in one of the studied kurgans."
    20. Gibbons. A.. 24 July 2015. Revolution in human evolution. Science. en. 349. 6246. 362–366. 10.1126/science.349.6246.362. 26206910. 2015Sci...349..362G. 0036-8075.
    21. Hanel. Andrea. Carlberg. Carsten. September 2020. Skin colour and vitamin D: An update. Experimental Dermatology. en. 29. 9. 864–875. 10.1111/exd.14142. 32621306. 0906-6705. free.
    22. Mathieson. Iain. Alpaslan-Roodenberg. Songül. Posth. Cosimo. Szécsényi-Nagy. Anna. Rohland. Nadin. Mallick. Swapan. Olalde. Iñigo. Broomandkhoshbacht. Nasreen. Candilio. Francesca. Cheronet. Olivia. Fernandes. Daniel. March 2018. The genomic history of southeastern Europe. Nature. en. 555. 7695. 197–203. 10.1038/nature25778. 1476-4687. 6091220. 29466330. 2018Natur.555..197M.
    23. Heyd. Volker. April 2017. Kossinna's smile . Antiquity. en. 91. 356. 348–359. 10.15184/aqy.2017.21. 0003-598X. 10138/255652. 164376362. free.
    24. Mathieson. Iain. Lazaridis. Iosif. Rohland. Nadin. Mallick. Swapan. Patterson. Nick. Roodenberg. Songül Alpaslan. Harney. Eadaoin. Stewardson. Kristin. Fernandes. Daniel. Novak. Mario. Sirak. Kendra. 24 December 2015. Genome-wide patterns of selection in 230 ancient Eurasians. Nature. en. 528. 7583. 499–503. 10.1038/nature16152. 0028-0836. 4918750. 26595274. 2015Natur.528..499M.
    25. Segurel . Laure . Why and when was lactase persistence selected for? Insights from Central Asian herders and ancient DNA . PLOS Biology . 2020 . 18 . 6 . e3000742 . 10.1371/journal.pbio.3000742 . 32511234 . 7302802 . free . "Furthermore, ancient DNA studies found that the LP mutation was absent or very rare in Europe until the end of the Bronze Age [26–29] and appeared first in individuals with steppe ancestry [19,20]. Thus, it was proposed that the mutation originated in Yamnaya-associated populations and arrived later in Europe by migration of these steppe herders."
    26. News: Callaway . Ewen . DNA data explosion lights up the Bronze Age . Nature. "the 101 sequenced individuals, the Yamnaya were most likely to have the DNA variation responsible for lactose tolerance, hinting that the steppe migrants might have eventually introduced the trait to Europe"
    27. Furholt . Martin . Massive Migrations? The Impact of Recent DNA Studies on our View of Third Millennium Europe . European Journal of Archaeology . 2018 . 21 . 2 . 159–191 . 10.1017/eaa.2017.43 . free . "For example, one lineage could have a biological evolutionary advantage over the other. Allentoft et al. (2015: 171) have found a remarkably high rate of lactose tolerance among individuals connected to Yamnaya and to Corded Ware, as opposed to the majority of Late Neolithic individuals."
    28. Saag . L . Human Genetics: Lactase Persistence in a Battlefield . Current Biology . 2020 . 30 . 21 . R1311–R1313 . 10.1016/j.cub.2020.08.087 . 33142099 . 226229587. free . 2020CBio...30R1311S .
    29. Book: Reich, David . Who we are and how we got here: ancient DNA and the new science of the human past. 2018. 978-0-19-882125-0 . Oxford, United Kingdom. 121. 1006478846.
    30. Lazaridis. Iosif . Alpaslan-Roodenberg . Songül . Acar . Ayşe . Açıkkol . Ayşen . Agelarakis . Anagnostis . Aghikyan . Levon . Akyüz . Uğur . Andreeva . Desislava . Andrijašević . Gojko . Antonović . Dragana . Armit . Ian . Atmaca . Alper . Avetisyan . Pavel . Aytek . Ahmet İhsan . Bacvarov . Krum . 26 August 2022 . The genetic history of the Southern Arc: A bridge between West Asia and Europe . Science . 377 . 6609 . eabm4247 . 10.1126/science.abm4247 . 36007055 . 10064553 . 251843620 . 0036-8075.
    31. Book: Reich, David . Who we are and how we got here: ancient DNA and the new science of the human past. 2018. 978-0-19-882125-0 . Oxford, United Kingdom. 239. 1006478846.
    32. Klejn . Leo . The Steppe Hypothesis of Indo-European Origins Remains to be Proven . Acta Archaeologica . 2017 . 88 . 1 . 193–204 . 10.1111/j.1600-0390.2017.12184.x.
    33. Balanovsky . O. . Genetic differentiation between upland and lowland populations shapes the Y-chromosomal landscape of West Asia . Human Genetics . 2017 . 136 . 4 . 437–450 . 10.1007/s00439-017-1770-2 . 28281087 . 3735168. "The ancient Yamnaya samples are located on the "eastern" R-GG400 branch of haplogroup R1b-L23, showing that the paternal descendants of the Yamnaya still live in the Pontic steppe and that the ancient Yamnaya population was not an important source of paternal lineages in present-day West Europeans."
    34. News: Zimmer . Carl . 10 June 2015 . DNA Deciphers Roots of Modern Europeans . New York Times . 12 December 2020.
    35. Book: Schneider . Thomas . Language Contact in Ancient Egypt . 2023 . LIT Verlag Münster . 978-3-643-91507-8 . 110-113 . en.
    36. Nikitin . Alexey . Ivanova . Svetlana . 28 November 2022 . Long-distance exchanges along the Black Sea coast in the Eneolithic and the steppe genetic ancestry problem . Cambridge Open Engage . en . 10.33774/coe-2022-7m315.
    37. Kroonen . Guus . Jakob . Anthony . Palmér . Axel I. . van Sluis . Paulus . Wigman . Andrew . 12 October 2022 . Indo-European cereal terminology suggests a Northwest Pontic homeland for the core Indo-European languages . PLOS ONE . 17 . 10 . e0275744 . 10.1371/journal.pone.0275744 . free . 1932-6203 . 9555676 . 36223379. 2022PLoSO..1775744K .
    38. Stoneking . Mark . Arias . Leonardo . Liu . Dang . Oliveira . Sandra . Pugach . Irina . Rodriguez . Jae Joseph Russell B. . 24 January 2023 . Genomic perspectives on human dispersals during the Holocene . Proceedings of the National Academy of Sciences . en . 120 . 4 . e2209475119 . 10.1073/pnas.2209475119 . 0027-8424 . 9942792 . 36649433. 2023PNAS..12009475S .
    39. Book: Kristiansen . Kristian . Kroonen . Guus . Willerslev . Eske . The Indo-European Puzzle Revisited: Integrating Archaeology, Genetics, and Linguistics . 11 May 2023 . Cambridge University Press . 978-1-009-26174-6 . 70-76 . en.
    40. Per, adding a north-Siberian people as a fourth reference population improves residuals for northeastern European populations. This accounts for the higher than expected Yamnaya contribution and brings it down to expected levels (67.8–50.4 % in Finns, 64.9–46.8 % in Russians).