Haplogroup R1a#R-M458 (R1a1a1b1a1)

The genetic divergence of R1a (M420) is estimated to have occurred 25,000{{sfn|Underhill et al.|2014}} years ago, which is the time of the last glacial maximum. A 2014 study by Peter A. Underhill et al., using 16,244 individuals from over 126 populations from across Eurasia, concluded that there was "a compelling case for the Middle East, possibly near present-day Iran, as the geographic origin of hg R1a".{{sfn|Underhill et al.|2014}} The ancient DNA record has shown the first R1a during the Mesolithic in Eastern Hunter-Gatherers (from Eastern Europe, c. 13,000 years ago),{{Cite journal|last1=Saag|first1=Lehti|last2=Vasilyev|first2=Sergey V.|last3=Varul|first3=Liivi|last4=Kosorukova|first4=Natalia V.|last5=Gerasimov|first5=Dmitri V.|last6=Oshibkina|first6=Svetlana V.|last7=Griffith|first7=Samuel J.|last8=Solnik|first8=Anu|last9=Saag|first9=Lauri|last10=D'Atanasio|first10=Eugenia|last11=Metspalu|first11=Ene|date=January 2021|title=Genetic ancestry changes in Stone to Bronze Age transition in the East European plain|url= |journal=Science Advances|language=en|volume=7|issue=4|pages=eabd6535|doi=10.1126/sciadv.abd6535|pmc=7817100|pmid=33523926|bibcode=2021SciA....7.6535S}}{{Cite journal|last1=Haak|first1=Wolfgang|last2=Lazaridis|first2=Iosif|last3=Patterson|first3=Nick|last4=Rohland|first4=Nadin|last5=Mallick|first5=Swapan|last6=Llamas|first6=Bastien|last7=Brandt|first7=Guido|last8=Nordenfelt|first8=Susanne|last9=Harney|first9=Eadaoin|last10=Stewardson|first10=Kristin|last11=Fu|first11=Qiaomei|date=February 10, 2015|title=Massive migration from the steppe is a source for Indo-European languages in Europe|url=https://www.biorxiv.org/content/10.1101/013433v1|journal=bioRxiv|language=en|pages=013433|doi=10.1101/013433|s2cid=196643946|arxiv=1502.02783|access-date=February 8, 2021|archive-date=December 23, 2019|archive-url=https://web.archive.org/web/20191223032847/https://www.biorxiv.org/content/10.1101/013433v1|url-status=live}} and the earliest case of R* among Upper Paleolithic Ancient North Eurasians,{{Cite journal|last1=Raghavan|first1=Maanasa|last2=Skoglund|first2=Pontus|last3=Graf|first3=Kelly E.|last4=Metspalu|first4=Mait|last5=Albrechtsen|first5=Anders|last6=Moltke|first6=Ida|last7=Rasmussen|first7=Simon|last8=Stafford Jr|first8=Thomas W.|last9=Orlando|first9=Ludovic|last10=Metspalu|first10=Ene|last11=Karmin|first11=Monika|date=January 2014|title=Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans|url= |journal=Nature|language=en|volume=505|issue=7481|pages=87–91|doi=10.1038/nature12736|pmc=4105016|pmid=24256729|bibcode=2014Natur.505...87R}} from which the Eastern Hunter-Gatherers predominantly derive their ancestry.{{Cite journal |last1=Narasimhan |first1=Vagheesh M. |last2=Patterson |first2=Nick|last3=Moorjani |first3=Priya|last4=Rohland |first4=Nadin |last5=Bernardos |first5=Rebecca |last6=Mallick |first6=Swapan |last7=Lazaridis |first7=Iosif |last8=Nakatsuka |first8=Nathan |last9=Olalde |first9=Iñigo |last10=Lipson |first10=Mark |last11=Kim |first11=Alexander M.|date=September 6, 2019|title=The formation of human populations in South and Central Asia |url= |journal=Science |language=en |volume=365 |issue=6457 |pages=eaat7487 |doi=10.1126/science.aat7487 |pmid=31488661 |pmc=6822619 |quote=Y chromosome haplogroup types R1b or R1a not represented in Iran and Turan in this period ...}} The genome of an individual belonging to the R1a5 subclade, dated to 10785–10626 BCE, from Peschanitsa, Arkhangelsk, Russia, and identified as a Western Russian Hunter-Gatherer, was published in January 2021.{{Cite journal |last=Saag |first=Lehti |last2=Vasilyev |first2=Sergey V. |last3=Varul |first3=Liivi |last4=Kosorukova |first4=Natalia V. |last5=Gerasimov |first5=Dmitri V. |last6=Oshibkina |first6=Svetlana V. |last7=Griffith |first7=Samuel J. |last8=Solnik |first8=Anu |last9=Saag |first9=Lauri |last10=D’Atanasio |first10=Eugenia |last11=Metspalu |first11=Ene |last12=Reidla |first12=Maere |last13=Rootsi |first13=Siiri |last14=Kivisild |first14=Toomas |last15=Scheib |first15=Christiana Lyn |date=January 20, 2021 |title=Genetic ancestry changes in Stone to Bronze Age transition in the East European plain |url=https://www.science.org/doi/10.1126/sciadv.abd6535 |journal=Science Advances |volume=7 |issue=4 |pages=eabd6535 |doi=10.1126/sciadv.abd6535 |pmc=7817100 |pmid=33523926}}

File:R1a origins (Underhill 2010) and R1a1a oldest expansion and highest frequency (2014).jpg

According to {{harvp|Underhill et al.|2014}}, the downstream M417 (R1a1a1) subclade diversified into Z282 (R1a1a1b1a) and Z93 (R1a1a1b2) circa 5,800 years ago "in the vicinity of Iran and Eastern Turkey".{{sfn|Underhill et al.|2014|p=130}}{{refn|group=note|According to Family Tree,{{who|reason=Source cited is not called that|date=July 2022}} they diversified c. 5,000 years ago.{{cite web |url=https://www.yfull.com/tree/R1a/ |title= R1a tree |website= YFull |access-date= July 15, 2016 |archive-date= August 19, 2016 |archive-url= https://web.archive.org/web/20160819051108/https://www.yfull.com/tree/R1a/ |url-status= live }}}} Even though R1a occurs as a Y-chromosome haplogroup among speakers of various languages such as Slavic and Indo-Iranian, the question of the origins of R1a1a is relevant to the Proto-Indo-European Urheimat hypotheses of the Proto-Indo-European people, and may also be relevant to the origins of the Indus Valley civilization. R1a shows a strong correlation with Indo-European languages of Southern and Western Asia, Central and Eastern Europe and to Scandinavia{{sfn|Mirabal et al.|2009}}{{sfn|Underhill et al.|2009}} being most prevalent in Eastern Europe, Central Asia, and South Asia. In Europe, Z282 is prevalent particularly while in Asia Z93 dominates. The connection between Y-DNA R-M17 and the spread of Indo-European languages was first noted by T. Zerjal and colleagues in 1999.{{cite book |first=T. |last=Zerjal |display-authors=et al |chapter=The use of Y-chromosomal DNA variation to investigate population history: recent male spread in Asia and Europe |editor1-first=S. S. |editor1-last=Papiha |editor2-first=R. |editor2-last=Deka |editor3-first=R. |editor3-last=Chakraborty |name-list-style=amp |title=Genomic diversity: applications in human population genetics |year=1999 |pages=91–101 |location=New York |publisher=Kluwer Academic/Plenum Publishers |isbn=978-0-3064-6295-5 |ref={{sfnref|Zerjal et al.|1999}}}}

{{See also|Indo-European migrations#Proto-Indo-Europeans|Indo-Aryan migration#Haplogroup R1a and related haplogroups}}

{{harvp|Semino et al.|2000}} proposed Ukrainian origins, and a postglacial spread of the R1a1 haplogroup during the Late Glacial Maximum, subsequently magnified by the expansion of the Kurgan culture into Europe and eastward.{{sfn|Semino et al.|2000}} Spencer Wells proposes Central Asian origins, suggesting that the distribution and age of R1a1 points to an ancient migration corresponding to the spread by the Kurgan people in their expansion from the Eurasian steppe.{{sfn|Wells|2001}} According to {{harvp|Pamjav et al.|2012}}, R1a1a diversified in the Eurasian Steppes or the Middle East and Caucasus region:

{{blockquote|Inner and Central Asia is an overlap zone for the R1a1-Z280 and R1a1-Z93 lineages [which] implies that an early differentiation zone of R1a1-M198 conceivably occurred somewhere within the Eurasian Steppes or the Middle East and Caucasus region as they lie between South Asia and Central- and Eastern Europe.{{sfn|Pamjav et al.|2012}}}}

Three genetic studies in 2015 gave support to the Kurgan theory of Gimbutas regarding the Indo-European Urheimat. According to those studies, haplogroups R1b and R1a, now the most common in Europe (R1a is also common in South Asia) would have expanded from the Pontic–Caspian steppes, along with the Indo-European languages; they also detected an autosomal component present in modern Europeans which was not present in Neolithic Europeans, which would have been introduced with paternal lineages R1b and R1a, as well as Indo-European languages.{{sfn|Haak et al.|2015}}{{sfn|Allentoft et al.|2015}}{{sfn|Mathieson et al.|2015}}

{{harvp|Silva et al.|2017}} noted that R1a in South Asia most "likely spread from a single Central Asian source pool, there do seem to be at least three and probably more R1a founder clades within the Indian subcontinent, consistent with multiple waves of arrival."{{sfn|Silva et al.|2017}} According to Martin P. Richards, co-author of {{harvp|Silva et al.|2017}}, the prevalence of R1a in India was "very powerful evidence for a substantial Bronze Age migration from central Asia that most likely brought Indo-European speakers to India."{{cite news |first=Tony |last=Joseph |date=June 16, 2017 |url=https://www.thehindu.com/sci-tech/science/how-genetics-is-settling-the-aryan-migration-debate/article19090301.ece |title=How genetics is settling the Aryan migration debate |work=The Hindu |access-date=June 2, 2019 |archive-date=October 4, 2023 |archive-url=https://web.archive.org/web/20231004150643/https://www.thehindu.com/sci-tech/science/how-genetics-is-settling-the-aryan-migration-debate/article19090301.ece |url-status=live }}{{Cite journal |last1=Silva |first1=Marina |last2=Oliveira |first2=Marisa |last3=Vieira |first3=Daniel |last4=Brandão |first4=Andreia |last5=Rito |first5=Teresa |last6=Pereira |first6=Joana B. |last7=Fraser |first7=Ross M. |last8=Hudson |first8=Bob |last9=Gandini |first9=Francesca |last10=Edwards |first10=Ceiridwen |last11=Pala |first11=Maria |last12=Koch |first12=John |last13=Wilson |first13=James F. |last14=Pereira |first14=Luísa |last15=Richards |first15=Martin B. |date=March 23, 2017 |title=A genetic chronology for the Indian Subcontinent points to heavily sex-biased dispersals |journal=BMC Evolutionary Biology |volume=17 |issue=1 |pages=88 |doi=10.1186/s12862-017-0936-9 |doi-access=free |issn=1471-2148 |pmc=5364613 |pmid=28335724|bibcode=2017BMCEE..17...88S }}

File:European-middle-neolithic-en.svg

File:Map Corded Ware culture-en.svg

David Anthony considers the Yamnaya culture to be the Indo-European Urheimat.{{sfn|Anthony|2007}}{{sfn|Anthony|Ringe|2015}} According to {{harvp|Haak et al.|2015}}, a massive migration from the Yamnaya culture northwards took place c. 2,500 BCE, accounting for 75% of the genetic ancestry of the Corded Ware culture, noting that R1a and R1b may have "spread into Europe from the East after 3,000 BCE".{{sfn|Haak et al.|2015|p=5}} Yet, all their seven Yamnaya samples belonged to the R1b-M269 subclade,{{sfn|Haak et al.|2015|p=5}} but no R1a1a has been found in their Yamnaya samples. This raises the question where the R1a1a in the Corded Ware culture came from, if it was not from the Yamnaya culture.{{sfn|Semenov|Bulat|2016}}

According to Marc Haber, the absence of haplogroup R1a-M458 in Afghanistan does not support a Pontic-Caspian steppe origin for the R1a lineages in modern Central Asian populations.{{harvnb|Haber et al.|2012|ps="R1a1a7-M458 was absent in Afghanistan, suggesting that R1a1a-M17 does not support, as previously thought [47], expansions from the Pontic Steppe [3], bringing the Indo-European languages to Central Asia and India."}}

According to Leo Klejn, the absence of haplogroup R1a in Yamnaya remains (despite its presence in Eneolithic Samara and Eastern Hunter Gatherer populations) makes it unlikely that Europeans inherited haplogroup R1a from Yamnaya.{{cite journal |last1=Klejn |first1=Leo S. |title=The Steppe Hypothesis of Indo-European Origins Remains to be Proven |journal=Acta Archaeologica |date=April 22, 2017 |volume=88 |issue=1 |pages=193–204 |doi=10.1111/j.1600-0390.2017.12184.x |url=https://brill.com/view/journals/acar/88/1/article-p193_13.xml |issn=0065-101X |access-date=November 23, 2022 |archive-date=December 25, 2022 |archive-url=https://web.archive.org/web/20221225050330/https://brill.com/view/journals/acar/88/1/article-p193_13.xml |url-status=live }} "As for the Y-chromosome, it was already noted in Haak, Lazaridis et al. (2015) that the Yamnaya from Samara had Y-chromosomes which belonged to R-M269 but did not belong to the clade common in Western Europe (p. 46 of supplement). Also, not a single R1a in Yamnaya unlike Corded Ware (R1a-dominated)."

Archaeologist Barry Cunliffe has said that the absence of haplogroup R1a in Yamnaya specimens is a major weakness in Haak's proposal that R1a has a Yamnaya origin.{{cite book |last1=Koch |first1=John T. |last2=Cunliffe |first2=Barry |title=Celtic from the West 3: Atlantic Europe in the Metal Ages |date=2016 |publisher=Oxbow Books |isbn=978-1-78570-228-0 |page=634 |url=https://books.google.com/books?id=Gv4sDwAAQBAJ&pg=PT634 |language=en |access-date=November 23, 2022 |archive-date=November 23, 2022 |archive-url=https://web.archive.org/web/20221123082348/https://books.google.com/books?id=Gv4sDwAAQBAJ&pg=PT634 |url-status=live }}

{{harvp|Semenov|Bulat|2016}} do argue for a Yamnaya origin of R1a1a in the Corded Ware culture, noting that several publications point to the presence of R1a1 in the Comb Ware culture.{{sfn|Semenov|Bulat|2016|p=41}}{{refn|group=note|{{harvp|Semenov|Bulat|2016}} refer to the following publications:{{ordered list|start=5

| {{cite journal|ref=none | biorxiv = 10.1101/013433 |doi=10.1038/NATURE14317 | bibcode=2015Natur.522..207H | title=Massive migration from the steppe is a source for Indo-European languages in Europe | year=2015 | last1 = Haak | first1 = Wolfgang | journal = Nature | volume = 522 | issue = 7555 | doi-access = free | pages = 207–211 | pmid = 25731166 | pmc = 5048219 | arxiv = 1502.02783}}

| {{cite bioRxiv|ref=none | biorxiv= 10.1101/016477 | title=Eight thousand years of natural selection in Europe | year=2015 | last1 = Mathieson | first1 = Iain }}

|item3-value=8 | Chekunova Е.М., Yartseva N.V., Chekunov М.К., Мazurkevich А.N. The First Results of the Genotyping of the Aboriginals and Human Bone Remains of the Archeological Memorials of the Upper Podvin'e. // Archeology of the lake settlements of IV—II Thousands BC: The chronology of cultures and natural environment and climatic rhythms. Proceedings of the International Conference, Devoted to the 50-year Research of the Pile Settlements on the North-West of Russia. St. Petersburg, November 13–15, 2014.

| {{cite journal|ref=none | pmid = 26567969 | doi=10.1038/ncomms9912 | volume=6 | title=Upper Palaeolithic genomes reveal deep roots of modern Eurasians | pmc=4660371 | year=2015 | journal=Nat Commun | page=8912 | last1 = Jones | first1 = ER | last2 = Gonzalez-Fortes | first2 = G | last3 = Connell | first3 = S | last4 = Siska | first4 = V | last5 = Eriksson | first5 = A | last6 = Martiniano | first6 = R | last7 = McLaughlin | first7 = RL | last8 = Gallego Llorente | first8 = M | last9 = Cassidy | first9 = LM | last10 = Gamba | first10 = C | last11 = Meshveliani | first11 = T | last12 = Bar-Yosef | first12 = O | last13 = Müller | first13 = W | last14 = Belfer-Cohen | first14 = A | last15 = Matskevich | first15 = Z | last16 = Jakeli | first16 = N | last17 = Higham | first17 = TF | last18 = Currat | first18 = M | last19 = Lordkipanidze | first19 = D | last20 = Hofreiter | first20 = M | last21 = Manica | first21 = A | last22 = Pinhasi | first22 = R | last23 = Bradley | first23 = DG| bibcode=2015NatCo...6.8912J }} }} }}

Kivisild et al. (2003) have proposed either South or West Asia,{{sfn|Kivisild et al.|2003}}{{refn|group=note|name="Kivisild2003"|Kivisild et al. (2003): "Haplogroup R1a, previously associated with the putative Indo-Aryan invasion, was found at its highest frequency in Punjab but also at a relatively high frequency (26%) in the Chenchu tribe. This finding, together with the higher R1a-associated short tandem repeat diversity in India and Iran compared with Europe and central Asia, suggests that southern and western Asia might be the source of this haplogroup."{{sfn|Kivisild et al.|2003}}}} while {{harvp|Mirabal et al.|2009}} see support for both South and Central Asia.{{sfn|Mirabal et al.|2009}} Sengupta et al. (2006) have proposed Indian origins.{{cite journal |ref={{harvid|Sengupta et al.|2006}} |vauthors=Sengupta S, Zhivotovsky LA, King R, Mehdi SQ, Edmonds CA, Chow CE, Lin AA, Mitra M, Sil SK, Ramesh A, Usha Rani MV, Thakur CM, Cavalli-Sforza LL, Majumder PP, Underhill PA |display-authors =6 |title=Polarity and temporality of high-resolution y-chromosome distributions in India identify both indigenous and exogenous expansions and reveal minor genetic influence of Central Asian pastoralists |journal=American Journal of Human Genetics |volume=78 |issue=2 |pages=202–221 |date=February 2006 |pmid=16400607 |pmc=1380230 |doi=10.1086/499411}}"Although considerable cultural impact on social hierarchy and language in South Asia is attributable to the arrival of nomadic Central Asian pastoralists, genetic data (mitochondrial and Y chromosomal) have yielded dramatically conflicting inferences on the genetic origins of tribes and castes of South Asia. We sought to resolve this conflict, using high-resolution data on 69 informative Y-chromosome binary markers and 10 microsatellite markers from a large set of geographically, socially, and linguistically representative ethnic groups of South Asia. We found that the influence of Central Asia on the pre-existing gene pool was minor. The ages of accumulated microsatellite variation in the majority of Indian haplogroups exceed 10,000–15,000 years, which attests to the antiquity of regional differentiation. Therefore, our data do not support models that invoke a pronounced recent genetic input from Central Asia to explain the observed genetic variation in South Asia. R1a1 and R2 haplogroups indicate demographic complexity that is inconsistent with a recent single history.ASSOCIATED MICROSATELLITE ANALYSES OF THE HIGH-FREQUENCY R1A1 HAPLOGROUP CHROMOSOMES INDICATE INDEPENDENT RECENT HISTORIES OF THE INDUS VALLEY AND THE PENINSULAR INDIAN REGION." Thanseem et al. (2006) have proposed either South or Central Asia.{{cite journal | vauthors = Thanseem I, Thangaraj K, Chaubey G, Singh VK, Bhaskar LV, Reddy BM, Reddy AG, Singh L | display-authors = 6 | title = Genetic affinities among the lower castes and tribal groups of India: inference from Y chromosome and mitochondrial DNA | journal = BMC Genetics | volume = 7 | pages = 42 | date = August 2006 | pmid = 16893451 | pmc = 1569435 | doi = 10.1186/1471-2156-7-42 | doi-access = free }} Sahoo et al. (2006) have proposed either South or West Asia.{{cite journal | vauthors = Sahoo S, Singh A, Himabindu G, Banerjee J, Sitalaximi T, Gaikwad S, Trivedi R, Endicott P, Kivisild T, Metspalu M, Villems R, Kashyap VK | display-authors = 6 | title = A prehistory of Indian Y chromosomes: evaluating demic diffusion scenarios | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 103 | issue = 4 | pages = 843–848 | date = January 2006 | pmid = 16415161 | pmc = 1347984 | doi = 10.1073/pnas.0507714103 | doi-access = free | bibcode = 2006PNAS..103..843S }} Thangaraj et al. (2010) have also proposed a South Asian origin.{{cite journal | vauthors = Thangaraj K, Naidu BP, Crivellaro F, Tamang R, Upadhyay S, Sharma VK, Reddy AG, Walimbe SR, Chaubey G, Kivisild T, Singh L | display-authors = 6 | title = The influence of natural barriers in shaping the genetic structure of Maharashtra populations | journal = PLOS ONE | volume = 5 | issue = 12 | pages = e15283 | date = December 2010 | pmid = 21187967 | pmc = 3004917 | doi = 10.1371/journal.pone.0015283 | veditors = Cordaux R | doi-access = free | bibcode = 2010PLoSO...515283T }} Sharma et al.(2009) theorizes the existence of R1a in India beyond 18,000 years to possibly 44,000 years in origin.{{sfn|Sharma et al.|2009}}

A number of studies from 2006 to 2010 concluded that South Asian populations have the highest STR diversity within R1a1a,{{sfn|Sengupta|2006}}{{sfn|Sahoo et al.|2006}}{{sfn|Mirabal et al.|2009}}{{sfn|Underhill et al.|2009}}{{sfn|Sharma et al.|2009}}{{sfn|Thangaraj et al.|2010}} and subsequent older TMRCA datings.{{refn|group=note|{{harvp|Sengupta|2006}}: "We found that the influence of Central Asia on the pre-existing gene pool was minor. The ages of accumulated microsatellite variation in the majority of Indian haplogroups exceed 10,000–15,000 years, which attests to the antiquity of regional differentiation. Therefore, our data do not support models that invoke a pronounced recent genetic input from Central Asia to explain the observed genetic variation in South Asia."}} R1a1a is present among both higher (Brahmin) castes and lower castes, and while the frequency is higher among Brahmin castes, the oldest TMRCA datings of the R1a haplogroup occur in the Saharia tribe, a scheduled caste of the Bundelkhand region of Central India.{{sfn|Sharma et al.|2009}}{{sfn|Thangaraj et al.|2010}}

From these findings some researchers argued that R1a1a originated in South Asia,{{sfn|Sahoo et al.|2006}}{{sfn|Sharma et al.|2009}}{{refn|group=note|name="R1a-India"|South-Asian origins:
* {{harvp|Sahoo et al.|2006}}: "... one should expect to observe dramatically lower genetic variation among Indian Rla lineages. In fact, the opposite is true: the STR haplotype diversity on the background of R1a in Central Asia (and also in Eastern Europe) has already been shown to be lower than that in India (6). Rather, the high incidence of R1* and Rla throughout Central Asian European populations (without R2 and R* in most cases) is more parsimoniously explained by gene flow in the opposite direction, possibly with an early founder effect in South or West Asia.{{sfn|Sahoo et al.|2006|p=845-846}}

* {{harvp|Sharma et al.|2009}}: "A peculiar observation of the highest frequency (up to 72.22%) of Y-haplogroup R1a1* in Brahmins hinted at its presence as a founder lineage for this caste group. Further, observation of R1a1* in different tribal population groups, existence of Y-haplogroup R1a* in ancestors and extended phylogenetic analyses of the pooled dataset of 530 Indians, 224 Pakistanis and 276 Central Asians and Eurasians bearing the R1a1* haplogroup supported the autochthonous origin of R1a1 lineage in India and a tribal link to Indian Brahmins. However, it is important to discover novel Y-chromosomal binary marker(s) for a higher resolution of R1a1* and confirm the present conclusions."}} excluding a more recent, yet minor, genetic influx from Indo-European migrants in northwestern regions such as Afghanistan, Balochistan, Punjab, and Kashmir.{{sfn|Sahoo et al.|2006}}{{sfn|Sengupta|2006}}{{sfn|Underhill et al.|2009}}{{refn|group=note|Though {{harvp|Sengupta|2006}} did concede that "[R1a1 and R2] could have actually arrived in southern India from a southwestern Asian source region multiple times." In full: "The widespread geographic distribution of HG R1a1-M17 across Eurasia and the current absence of informative subdivisions defined by binary markers leave uncertain the geographic origin of HG R1a1-M17. However, the contour map of R1a1-M17 variance shows the highest variance in the northwestern region of India ... The question remains of how distinctive is the history of L1 relative to some or all of R1a1 and R2 representatives. This uncertainty neutralizes previous conclusions that the intrusion of HGs R1a1 and R2 from the northwest in Dravidian-speaking southern tribes is attributable to a single recent event. [R1a1 and R2] could have actually arrived in southern India from a southwestern Asian source region multiple times, with some episodes considerably earlier than others. Considerable archeological evidence exists regarding the presence of Mesolithic peoples in India (Kennedy 2000), some of whom could have entered the subcontinent from the northwest during the late Pleistocene epoch. The high variance of R1a1 in India (table 12), the spatial frequency distribution of R1a1 microsatellite variance clines (fig. 4), and expansion time (table 11) support this view."{{sfn|Sengupta|2006}}}}

The conclusion that R1a originated in India has been questioned by more recent research,{{sfn|Silva et al.|2017}}{{cite book | last=Lalueza-Fox | first=C. | title=Inequality: A Genetic History | publisher=MIT Press | year=2022 | isbn=978-0-262-04678-7 | url=https://books.google.com/books?id=xLZNEAAAQBAJ&pg=PA81 | access-date=July 16, 2023 | pages=81–82 | archive-date=July 16, 2023 | archive-url=https://web.archive.org/web/20230716102536/https://books.google.com/books?id=xLZNEAAAQBAJ&pg=PA81 | url-status=live }}{{refn|group=note|Lalueza-Fox: "Some years ago, local scientists supported the view that the existence of an R1a Y chromosome was not attributable to a foreign gene flow but instead that this lineage had emerged on the subcontinent and spread from there. But the phylogenetic reconstruction of this haplogroup did not support this view."}} offering proof that R1a arrived in India with multiple waves of migration.{{sfn|Silva et al.|2017}}{{sfn|Narasimhan et al.|2019}}

{{See also|Kura–Araxes culture|Uruk period|Indus Valley Civilisation#Possible near-Eastern Dravidian origins}}

{{harvp|Haak et al.|2015}} found that part of the Yamnaya ancestry derived from the Middle East and that neolithic techniques probably arrived at the Yamnaya culture from the Balkans.{{refn|group=note|Yet, Haak et al. also explicitly state: "a type of Near Eastern ancestry different from that which was introduced by early farmers".{{clarify|reason=This "statement" has no verb, it is merely a noun phrase with no factual assertion|date=July 2022}}{{sfn|Haak et al.|2015|p=4}}}} The Rössen culture (4,600–4,300 BC), which was situated on Germany and predates the Corded Ware culture, an old subclade of R1a, namely L664, can still be found.{{refn|group=note|According to Family Tree DNA, L664 formed 4,700 ybp, that is, 2,700 BCE.}}

Part of the South Asian genetic ancestry derives from west Eurasian populations, and some researchers have implied that Z93 may have come to India via Iran{{sfn|Mascarenhas et al.|2015|p=9}} and expanded there during the Indus Valley civilization.{{sfn|Underhill et al.|2014}}{{sfn|Poznik et al.|2016|p=5}}

{{harvp|Mascarenhas et al.|2015}} proposed that the roots of Z93 lie in West Asia, and proposed that "Z93 and L342.2 expanded in a southeasterly direction from Transcaucasia into South Asia",{{sfn|Mascarenhas et al.|2015|p=9}} noting that such an expansion is compatible with "the archeological records of eastward expansion of West Asian populations in the 4th millennium BCE culminating in the so-called Kura-Araxes migrations in the post-Uruk IV period."{{sfn|Mascarenhas et al.|2015|p=9}} Yet, Lazaridis noted that sample I1635 of {{harvp|Lazaridis et al.|2016}}, their Armenian Kura-Araxes sample, carried Y-haplogroup R1b1-M415(xM269){{refn|group=note|Lazaridis, [https://twitter.com/iosif_lazaridis/status/744192603424456704 Twitter, 18 June 2016]: "I1635 (Armenia_EBA) is R1b1-M415(xM269). We'll be sure to include in the revision. Thanks to the person who noticed! #ILovePreprints."{{Unreliable source?|reason=Twitter is generally not acceptable as a source|date=July 2022}}
See also {{cite web |website=Eurogenes Blog |date=November 26, 2016 |url= http://eurogenes.blogspot.nl/2016/11/big-deal-of-2016-territory-of-present.html |title=Big deal of 2016: the territory of present-day Iran cannot be the Indo-European homeland |postscript=,}}{{Unreliable source?|reason=Blogs are generally not acceptable as sources|date=July 2022}} for a discussion of the same topic.}} (also called R1b1a1b-CTS3187).Arame's English blog, [http://aramepal.blogspot.nl/2016_11_01_archive.html Y DNA from ancient Near East] {{Webarchive|url=https://web.archive.org/web/20161127214951/http://aramepal.blogspot.nl/2016_11_01_archive.html |date=November 27, 2016 }}{{Unreliable source?|reason=Blogs are generally not acceptable as sources|date=July 2022}}

According to {{harvp|Underhill et al.|2014}} the diversification of Z93 and the "early urbanization within the Indus Valley ... occurred at [5,600 years ago] and the geographic distribution of R1a-M780 (Figure 3d{{refn|group=note|See map for M780 distribution at Dieneke's Anthropology Blog, Major new article on the deep origins of Y-haplogroup R1a (Underhill et al. 2014){{Cite web|url=http://dienekes.blogspot.com/2014/03/major-new-article-on-deep-origins-of-y.html|title=Dienekes' Anthropology Blog: Major new article on the deep origins of Y-haplogroup R1a (Underhill et al. 2014)|date=March 27, 2014|access-date=December 20, 2019|archive-date=December 20, 2019|archive-url=https://web.archive.org/web/20191220175749/http://dienekes.blogspot.com/2014/03/major-new-article-on-deep-origins-of-y.html|url-status=live}}{{Unreliable source?|reason=Blogs are generally not acceptable as sources|date=July 2022}}}}) may reflect this."{{sfn|Underhill et al.|2014}}{{refn|group=note|name="FamilyTree-M780"|According to Family Tree DNA, M780 formed 4700 ybp. This dating coincides with the eastward movement between 2800 and 2600 BCE of the Yamnaya culture into the region of the Poltavka culture, a predecessor of the Sintashta culture, from which the Indo-Iranians originated. M780 is concentrated in the Ganges Valley, the locus of the classic Vedic society.}} {{harvp|Poznik et al.|2016}} note that "striking expansions" occurred within R1a-Z93 at c. 4,500–4,000 years ago, which "predates by a few centuries the collapse of the Indus Valley Civilisation."{{sfn|Poznik et al.|2016|p=5}}{{refn|group=note|{{harvp|Poznik et al.|2016}} calculate with a generation time of 30 years; a generation time of 20 years yields other results.}}

However, according to {{harvp|Narasimhan et al.|2018}}, steppe pastoralists are a likely source for R1a in India.{{sfn|Narasimhan et al.|2018}}{{refn|group=note|1="The evidence that the Steppe_MLBA [Middle to Late Bronze Age] cluster is a plausible source for the Steppe ancestry in South Asia is also supported by Y chromosome evidence, as haplogroup R1a which is of the Z93 subtype common in South Asia today [Underhill et al. (2014), Silva et al. (2017)] was of high frequency in Steppe_MLBA (68%) (16), but rare in Steppe_EMBA [Early to Middle Bronze Age] (absent in our data)."{{sfn|Narasimhan et al.|2018}}}}

The R1a family tree now has three major levels of branching, with the largest number of defined subclades within the dominant and best known branch, R1a1a (which will be found with various names such as "R1a1" in relatively recent but not the latest literature).

The topology of R1a is as follows (codes [in brackets] non-isogg codes):{{Cite web|url=https://www.familytreedna.com/groups/r-1a/about/results|title=About Us|website=Family Tree DNA|access-date=December 20, 2019|archive-date=August 15, 2019|archive-url=https://web.archive.org/web/20190815072247/https://www.familytreedna.com/groups/r-1a/about/results|url-status=live}}{{verify source|date=July 2020|reason=Did not find info on "About Us" page.}}{{Cite web |url=https://isogg.org/tree/ISOGG_HapgrpR.html |title=ISOGG 2017 Y-DNA Haplogroup R |website=isogg.org |access-date=December 20, 2019 |archive-date=February 10, 2007 |archive-url=https://web.archive.org/web/20070210011401/https://isogg.org/tree/ISOGG_HapgrpR.html |url-status=live }}{{sfn|Underhill et al.|2014}}{{Cite web|url=https://www.snpedia.com/index.php/Haplogroup_R_(Y-DNA)|title=Haplogroup R (Y-DNA) - SNPedia|website=www.snpedia.com|access-date=December 20, 2019|archive-date=May 5, 2018|archive-url=https://web.archive.org/web/20180505070347/https://www.snpedia.com/index.php/Haplogroup_R_(Y-DNA)|url-status=live}} Tatiana et al. (2014) "rapid diversification process of K-M526 likely occurred in Southeast Asia, with subsequent westward expansions of the ancestors of haplogroups R and Q."{{sfn|Karafet et al.|2014}}

{{Tree list}}

• P P295/PF5866/S8 (also known as K2b2).
• {{Tree list/final branch}}R (R-M207)
• R*
• R1 (R-M173)
• R1*
• R1a (M420) (Eastern Europe, Asia){{sfn|Underhill et al.|2014}}
• R1a*
• {{Tree list/final branch}}R1a1 (M459/PF6235, SRY1532.2/SRY10831.2)
• R1a1 (M459)
• {{Tree list/final branch}}R1a1a (M17, M198)
• {{Tree list/final branch}}R1a1a1 (M417, page7)
• R1a1a1a (CTS7083/L664/S298)
• R1a1a1b (S224/Z645, S441/Z647)
• R1a1a1b1 (PF6217/S339/Z283)
• {{Tree list/final branch}}R1a1a1b1a (Z282) [R1a1a1a*] (Z282) {{sfn|Underhill et al.|2014|p=125}} (Eastern Europe)
• R1a1a1b1a1 [The old topological code is R1a1a1b*，which is outdated and might lead to some confusion.]{{sfn|Underhill et al.|2014|p=125}} (M458){{sfn|Underhill et al.|2014|p=125}} [R1a1a1g] (M458)
• [R1a1a1g*]
• [R1a1a1g1] (M334)
• {{Tree list/final branch}}R1a1a1b1a1a (L260/S222) [R1a1a1g2]
• R1a1a1b1a2 (S466/Z280, S204/Z91)
• R1a1a1b1a2a
• {{Tree list/final branch}}R1a1a1b1a2b (CTS1211) [R1a1a1c*] (M558){{sfn|Underhill et al.|2014|p=125}} [R-CTS1211] (V2803/CTS3607/S3363/M558, CTS1211/S3357, Y34/FGC36457)
• R1a1a1b1a2b3* (M417+, Z645+, Z283+, Z282+, Z280+, CTS1211+, CTS3402, Y33+, CTS3318+, Y2613+) (Gwozdz's Cluster K){{verify source|date=July 2020|reason=Did not find info on "About Us" page.}}
• {{Tree list/final branch}}R1a1a1b1a2b3a (L365/S468)
• {{Tree list/final branch}}R1a1a1b1a3 (Z284) [R1a1a1a1] (Z284){{sfn|Underhill et al.|2014|p=125}}
• {{Tree list/final branch}}R1a1a1b2 (F992/S202/Z93) [R1a1a2*] (Z93, M746){{sfn|Underhill et al.|2014|p=125}} (Central Asia, South Asia and West Asia)
• {{Tree list/final branch}}R1a1a1b2a (F3105/S340/Z94, L342.2/S278.2) [R1a1b2a*] (Z95){{sfn|Underhill et al.|2014|p=125}} R-Z94 (Z94/F3105/S340, Z95/F3568)
• R-Z2124 (Z2121/S3410, Z2124)
• {{Tree list/final branch}}[R1a1b2a*] (Z2125){{sfn|Underhill et al.|2014|p=125}}
• [R1a1b2a*] (M434){{sfn|Underhill et al.|2014|p=125}} [R1a1a1f] (M434)
• {{Tree list/final branch}}[R1a1b2a*] (M204){{sfn|Underhill et al.|2014|p=125}}
• [R1a1b2a1*] (M560){{sfn|Underhill et al.|2014|p=125}}
• [R1a1b2a2*] (M780, L657){{sfn|Underhill et al.|2014|p=125}} (India){{sfn|Underhill et al.|2014}}
• {{Tree list/final branch}}[R1a1b2a3*] (Z2122, M582){{sfn|Underhill et al.|2014|p=125}}
• [R1a1a1c] (M64.2, M87, M204)
• [R1a1a1d] (P98)
• {{Tree list/final branch}}[R1a1a1d2a]{{Cite web |website=Eurogenes Blog |date=March 21, 2016 |url= http://eurogenes.blogspot.no/2016/03/r1a-in-yamnaya.html |title=R1a in Yamnaya |access-date=December 20, 2019 |archive-url= https://web.archive.org/web/20180505065717/http://eurogenes.blogspot.no/2016/03/r1a-in-yamnaya.html |archive-date=May 5, 2018 |url-status=dead}}
• {{Tree list/final branch}}[R1a1a1e] (PK5)
• {{Tree list/final branch}}R1b (M343) (Western Europe)
• {{Tree list/final branch}}R2 (India)

{{Tree list/end}}

R1a is distinguished by several unique markers, including the M420 mutation. It is a subclade of Haplogroup R-M173 (previously called R1). R1a has the sister-subclades Haplogroup R1b-M343, and the paragroup R-M173*.

R1a, defined by the mutation M420, has two primary branches: R-M459 (R1a1) and R-YP4141 (R1a2).

As of 2025, ten ancient basal R1a* genotypes have been recovered and published, from remains found in Estonia, Poland, Russia, and Ukraine; the oldest sample (Vasilevka 497) dated to c. 8700 BCE, and excavated in the Vasylivka, Bakhmut Raion, Donetsk Oblast.{{Cite web |title=haplotree.info - ancientdna.info. Map based on All Ancient DNA v. 2.07.26. |url=https://haplotree.info/maps/ancient_dna/slideshow_samples.php?searchcolumn=Y_Haplotree_Variant&searchfor=R-M459*&ybp=500000,0 |access-date=January 19, 2025 |website=haplotree.info}}

R1a2 (R-YP4141) has two branches R1a2a (R-YP5018) and R1a2b (R-YP4132).{{cite web | url=https://www.yfull.com/arch-5.07/tree/R1a/ | title=R1a YTree }}

This rare primary subclade was initially regarded as part of a paragroup of R1a*, defined by SRY1532.2 (and understood to always exclude M459 and its synonyms SRY10831.2, M448, L122, and M516).{{sfn|Underhill et al.|2009}}{{cite web |last1=Krahn |first1=Thomas |title=Draft Y-Chromosome Tree |website=Family Tree DNA |url=http://ytree.ftdna.com/index.php?name=Draft&parent=99812767 |access-date=December 7, 2012 |archive-url=https://web.archive.org/web/20130526205543/http://ytree.ftdna.com/index.php?name=Draft&parent=99812767 |archive-date=May 26, 2013 |url-status=dead }}

YP4141 later replaced SRY1532.2 – which was found to be unreliable – and the R1a(xR-M459) group was redefined as R1a2. It is relatively unusual, though it has been tested in more than one survey. {{harvp|Sahoo et al.|2006}} reported R-SRY1532.2* for 1/15 Himachal Pradesh Rajput samples.{{sfn|Sahoo et al.|2006}} Underhill et al. (2009) reported 1/51 in Norway, 3/305 in Sweden, 1/57 Greek Macedonians, 1/150 (or 2/150) Iranians, 2/734 ethnic Armenians, 1/141 Kabardians, 1/121 Omanis, 1/164 in the United Arab Emirates, and 3/612 in Turkey. Testing of 7224 more males in 73 other Eurasian populations showed no sign of this category.{{sfn|Underhill et al.|2009}}

The oldest known example genotyped is from a set of remains, dating to c. 3500 BCE, recovered from the Kumyshanskaya Cave, in Russia.

The major subclade R-M459 includes an overwhelming majority of individuals within R1a more broadly.

Ancient R-M459 genotypes, dating to c. 8650 BCE, have been recovered from two sets of remains excavated at Minino, Russia.

R-YP1272, also known as R-M459(xM198), is an extremely rare primary subclade of R1a1. It has been found in three individuals, from Belarus, Tunisia and the Coptic community in Egypt respectively.{{cite web | url=https://www.yfull.com/arch-5.07/tree/R-M459/ | title=R-M459 YTree }}

The following SNPs are associated with R1a1a:

R1a1a1 (R-M417) is the most widely found subclade, in two variations which are found respectively in Europe (R1a1a1b1 (R-Z282) ([R1a1a1a*] (R-Z282) (Underhill 2014){{sfn|Underhill et al.|2014}}) and Central and South Asia (R1a1a1b2 (R-Z93) ([R1a1a2*] (R-Z93) Underhill 2014){{sfn|Underhill et al.|2014}}).

The oldest known basal R1a1a1 genotype so far published has been dated to c. 5650 BCE, and was recovered from a site at Trestiana, Romania.

This large subclade appears to encompass most of the R1a1a found in Europe.{{sfn|Pamjav et al.|2012}}

• R1a1a1b1a [R1a1a1a* (Underhill (2014))] (R-Z282*) occurs in northern Ukraine, Belarus, and Russia at a frequency of c. 20%.{{sfn|Underhill et al.|2014}}
• R1a1a1b1a3 [R1a1a1a1 (Underhill (2014))] (R-Z284) occurs in Northwest Europe and peaks at c. 20% in Norway.{{sfn|Underhill et al.|2014}}
• R1a1a1c (M64.2, M87, M204) is apparently rare: it was found in 1 of 117 males typed in southern Iran.{{sfn|Regueiro|2006}}

File:R-M458 frequency distribution.PNG

R-M458 is a mainly Slavic SNP, characterized by its own mutation, and was first called cluster N. Underhill et al. (2009) found it to be present in modern European populations roughly between the Rhine catchment and the Ural Mountains and traced it to "a founder effect that ... falls into the early Holocene period, 7.9±2.6 KYA." (Zhivotovsky speeds, 3x overvalued){{sfn|Underhill et al.|2009}} M458 was found in one skeleton from a 14th-century grave field in Usedom, Mecklenburg-Vorpommern, Germany.{{cite thesis |last1=Freder |first1=Janine |trans-title=Anthropological investigation in due consideration of the ethnical background |title=Die mittelalterlichen Skelette von Usedom: Anthropologische Bearbeitung unter besonderer Berücksichtigung des ethnischen Hintergrundes |language=de |date=2010 |doi=10.17169/refubium-8995 |page=86 |publisher=Freie Universität Berlin }} The paper by Underhill et al. (2009) also reports a surprisingly high frequency of M458 in some Northern Caucasian populations (18% among Ak Nogai,https://cyberleninka.ru/article/n/tyurki-kavkaza-sravnitelnyy-analiz-genofondov-po-dannym-o-y-hromosome {{Webarchive|url=https://web.archive.org/web/20231107210034/https://cyberleninka.ru/article/n/tyurki-kavkaza-sravnitelnyy-analiz-genofondov-po-dannym-o-y-hromosome |date=November 7, 2023 }} "высокая частота R1a среди кубанских ногайцев (субветвь R1a1a1g-M458 забирает 18%" 7.8% among Qara Nogai and 3.4% among Abazas).{{cite journal | pmc=2987245 | date=2009 | last1=Underhill | first1=P. A. | last2=Myres | first2=N. M. | last3=Rootsi | first3=S. | last4=Metspalu | first4=M. | last5=Zhivotovsky | first5=L. A. | last6=King | first6=R. J. | last7=Lin | first7=A. A. | last8=Chow | first8=C. E. | last9=Semino | first9=O. | last10=Battaglia | first10=V. | last11=Kutuev | first11=I. | last12=Järve | first12=M. | last13=Chaubey | first13=G. | last14=Ayub | first14=Q. | last15=Mohyuddin | first15=A. | last16=Mehdi | first16=S. Q. | last17=Sengupta | first17=S. | last18=Rogaev | first18=E. I. | last19=Khusnutdinova | first19=E. K. | last20=Pshenichnov | first20=A. | last21=Balanovsky | first21=O. | last22=Balanovska | first22=E. | last23=Jeran | first23=N. | last24=Augustin | first24=D. H. | last25=Baldovic | first25=M. | last26=Herrera | first26=R. J. | last27=Thangaraj | first27=K. | last28=Singh | first28=V. | last29=Singh | first29=L. | last30=Majumder | first30=P. | title=Separating the post-Glacial coancestry of European and Asian y chromosomes within haplogroup R1a | journal=European Journal of Human Genetics | volume=18 | issue=4 | pages=479–484 | doi=10.1038/ejhg.2009.194 | pmid=19888303 | display-authors=1 }}

R1a1a1b1a1a (R-L260), commonly referred to as West Slavic or Polish, is a subclade of the larger parent group R-M458, and was first identified as an STR cluster by {{harvnb|Pawlowski et al.|2002}}. In 2010 it was verified to be a haplogroup identified by its own mutation (SNP).{{cite web |first=Peter |last=Gwozdz |date=August 6, 2018 |url=http://www.gwozdz.org/PolishClades.html#L260M458News |title=Polish Y-DNA Clades |access-date=July 15, 2016 |archive-date=July 15, 2016 |archive-url=https://web.archive.org/web/20160715055534/http://www.gwozdz.org/PolishClades.html#L260M458News |url-status=live }} It apparently accounts for about 8% of Polish men, making it the most common subclade in Poland. Outside of Poland it is less common.{{sfn|Pawlowski et al.|2002}} In addition to Poland, it is mainly found in the Czech Republic and Slovakia, and is considered "clearly West Slavic". The founding ancestor of R-L260 is estimated to have lived between 2000 and 3000 years ago, i.e. during the Iron Age, with significant population expansion less than 1,500 years ago.{{sfn|Gwozdz|2009}}

R-M334 ([R1a1a1g1], a subclade of [R1a1a1g] (M458) c.q. R1a1a1b1a1 (M458)) was found by Underhill et al. (2009) only in one Estonian man and may define a very recently founded and small clade.{{sfn|Underhill et al.|2009}}

R1a1a1b1a2b3* (M417+, Z645+, Z283+, Z282+, Z280+, CTS1211+, CTS3402, Y33+, CTS3318+, Y2613+) (Gwozdz's Cluster K){{verify source|date=July 2020|reason=Did not find info on "About Us" page.}} is a STR based group that is R-M17(xM458). This cluster is common in Poland but not exclusive to Poland.{{sfn|Gwozdz|2009}}

R1a1a1b1a2b3a (R-L365) was early called Cluster G.{{citation needed|date=July 2016}}

This large subclade appears to encompass most of the R1a1a found in Asia, being related to Indo-European migrations (including Scythians, Indo-Aryan migrations and so on).{{sfn|Pamjav et al.|2012}}

• R-Z93* or R1a1a1b2* (R1a1a2* in Underhill (2014)) is most common (>30%) in the South Siberian Altai region of Russia, cropping up in Kyrgyzstan (6%) and in all Iranian populations (1-8%).{{sfn|Underhill et al.|2014}} The oldest published R-Z93 genotypes being sampled from graves, dated to c. 2650 - 2700 BCE, in Naumovskoye, and Khanevo, Vologda Oblast, and Khaldeevo, Rostov District, Russia.
• R-Z2125 occurs at highest frequencies in Kyrgyzstan and in Afghan Pashtuns (>40%). At a frequency of >10%, it is also observed in other Afghan ethnic groups and in some populations in the Caucasus and Iran.{{sfn|Underhill et al.|2014}}
• R-M434 (R1a1a6) is a subclade of Z2125. It was detected in 14 people (out of 3667 people tested), all in a restricted geographical range from Pakistan to Oman. This likely reflects a recent mutation event in Pakistan.{{sfn|Underhill et al.|2009}}
• R-M560 is very rare and was only observed in four samples: two Burushaski speakers (north Pakistan), one Hazara (Afghanistan), and one Iranian Azerbaijani.{{sfn|Underhill et al.|2014}}
• R-M780 (R1a1b2a2*) occurs at high frequency in South Asia: India, Pakistan, Afghanistan, and the Himalayas. Turkey share R1a (12.1%) sublineages.{{cite journal | pmc=8433500 | year=2021 | last1=Kars | first1=M. E. | last2=Başak | first2=A. N. | last3=Onat | first3=O. E. | last4=Bilguvar | first4=K. | last5=Choi | first5=J. | last6=Itan | first6=Y. | last7=Çağlar | first7=C. | last8=Palvadeau | first8=R. | last9=Casanova | first9=J. L. | last10=Cooper | first10=D. N. | last11=Stenson | first11=P. D. | last12=Yavuz | first12=A. | last13=Buluş | first13=H. | last14=Günel | first14=M. | last15=Friedman | first15=J. M. | last16=Özçelik | first16=T. | title=The genetic structure of the Turkish population reveals high levels of variation and admixture | journal=Proceedings of the National Academy of Sciences of the United States of America | volume=118 | issue=36 | pages=e2026076118 | doi=10.1073/pnas.2026076118 | pmid=34426522 | bibcode=2021PNAS..11826076K | doi-access=free }} Roma from Slovakia share 3% of R1a{{cite journal |doi=10.1537/ase.080422 |title=Y-haplogroup frequencies in the Slovak Romany population |year=2009 |last1=Petrejcíková |first1=EVA |last2=Soták |first2=Miroslav |last3=Bernasovská |first3=Jarmila |last4=Bernasovský |first4=Ivan |last5=Sovicová |first5=Adriana |last6=Bôziková |first6=Alexandra |last7=Boronová |first7=Iveta |last8=Švícková |first8=Petra |last9=Gabriková |first9=Dana |last10=MacEková |first10=Sona |journal=Anthropological Science |volume=117 |issue=2 |pages=89–94 |doi-access=free }} The group also occurs at >3% in some Iranian populations and is present at >30% in Roma from Croatia and Hungary.{{sfn|Underhill et al.|2014}}

File:Y-Haplogroup R1 distribution.png

In Mesolithic Europe, R1a is characteristic of Eastern Hunter-Gatherers (EHGs).{{sfn|Saag et al.|2020|p=5}} A male EHG of the Veretye culture buried at Peschanitsa near Lake Lacha in Arkhangelsk Oblast, Russia c. 10,700 BCE was found to be a carrier of the paternal haplogroup R1a5-YP1301 and the maternal haplogroup U4a.{{sfn|Saag et al.|2020|p=29|loc=Table 1}}{{sfn|Saag et al.|2020|loc=Supplementary Data 2, Row 4}}{{sfn|Saag et al.|2020|p=5}} A male, named PES001, from Peschanitsa in northwestern Russia was found to carry R1a5, and dates to at least 10,600 years ago. More examples include the males Minino II (V) and Minino II (I/1), with the former carrying R1a1 and the latter R1a respectively, with the former being at 10,600 years old and the latter at least 10,400 years old respectively, both from Minino in northwestern Russia.{{Cite journal |last1=Posth |first1=Cosimo |last2=Yu |first2=He |last3=Ghalichi |first3=Ayshin |last4=Rougier |first4=Hélène |last5=Crevecoeur |first5=Isabelle |last6=Huang |first6=Yilei |last7=Ringbauer |first7=Harald |last8=Rohrlach |first8=Adam B. |last9=Nägele |first9=Kathrin |last10=Villalba-Mouco |first10=Vanessa |last11=Radzeviciute |first11=Rita |last12=Ferraz |first12=Tiago |last13=Stoessel |first13=Alexander |last14=Tukhbatova |first14=Rezeda |last15=Drucker |first15=Dorothée G. |date=March 1, 2023 |title=Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers |journal=Nature |language=en |volume=615 |issue=7950 |pages=117–126 |doi=10.1038/s41586-023-05726-0 |pmid=36859578 |pmc=9977688 |bibcode=2023Natur.615..117P |issn=1476-4687|hdl=10256/23099 |hdl-access=free }} A Mesolithic male from Karelia c. 8,800 BCE to 7950 BCE has been found to be carrying haplogroup R1a.{{sfn|Fu et al.|2016}} A Mesolithic male buried at Deriivka c. 7000 BCE to 6700 BCE carried the paternal haplogroup R1a and the maternal U5a2a.{{sfn|Mathieson et al.|2015}} Another male from Karelia from c. 5,500 to 5,000 BC, who was considered an EHG, carried haplogroup R1a.{{sfn|Haak et al.|2015}} A male from the Comb Ceramic culture in Kudruküla c. 5,900 BCE to 3,800 BCE has been determined to be a carrier of R1a and the maternal U2e1.{{sfn|Saag et al.|2017}} According to archaeologist David Anthony, the paternal R1a-Z93 was found at the Oskol river near a no longer existing kolkhoz "Alexandria", Ukraine c. 4000 BCE, "the earliest known sample to show the genetic adaptation to lactase persistence (13910-T)."{{sfn|Anthony|2019|pp=16, 17}} R1a has been found in the Corded Ware culture,{{sfn|Haak et al.|2008}}{{sfn|Brandit et al.|2013}} in which it is predominant.{{sfn|Malmström et al.|2019|p=2}} Examined males of the Bronze Age Fatyanovo culture belong entirely to R1a, specifically subclade R1a-Z93.{{sfn|Saag et al.|2020|p=5}}{{sfn|Saag et al.|2020|p=29|loc=Table 1}}{{sfn|Saag et al.|2020|loc=Supplementary Data 2, Rows 5-49}}

Haplogroup R1a has later been found in ancient fossils associated with the Urnfield culture;{{cite web |url=http://dirkschweitzer.net/LichtensteinCaveAnalysis0804DS.pdf |last=Schweitzer |first=D. |title=Lichtenstein Cave Data Analysis |publisher=dirkschweitzer.net |date=March 23, 2008 |url-status=dead |archive-url=https://web.archive.org/web/20110814164431/http://dirkschweitzer.net/LichtensteinCaveAnalysis0804DS.pdf |archive-date=August 14, 2011}} Summary in English of {{harvp|Schilz|2006}}. as well as the burial of the remains of the Sintashta,{{sfn|Allentoft et al.|2015}} Andronovo,{{sfn|Keyser et al.|2009}} the Pazyryk,{{sfn|Ricaut et al.|2004}} Tagar,{{sfn|Keyser et al.|2009}} Tashtyk,{{sfn|Keyser et al.|2009}} and Srubnaya cultures, the inhabitants of ancient Tanais,Korniyenko, I. V.; Vodolazhsky D. I. {{lang|ru|italic=no|cat=no|"Использование нерекомбинантных маркеров Y-хромосомы в исследованиях древних популяций (на примере поселения Танаис)"}} [The use of non-recombinant markers of the Y-chromosome in the study of ancient populations (on the example of the settlement of Tanais)]. {{lang|ru|italic=no|cat=no|Материалы Донских антропологических чтений}} [Materials of the Don Anthropological Readings]. Rostov-on-Don: Rostov Research Institute of Oncology, 2013. in the Tarim mummies,{{sfn|Chunxiang Li et al.|2010}} and the aristocracy of Xiongnu.{{sfn|Kim et al.|2010}} The skeletal remains of a father and his two sons, from an archaeological site discovered in 2005 near Eulau (in Saxony-Anhalt, Germany) and dated to about 2600 BCE, tested positive for the Y-SNP marker SRY10831.2. The Ysearch number for the Eulau remains is 2C46S. The ancestral clade was thus present in Europe at least 4600 years ago, in association with one site of the widespread Corded Ware culture.{{sfn|Haak et al.|2008}}

In Europe, the R1a1a sub-clade is primarily characteristic of Balto-Slavic populations, with two exceptions: southern Slavs and northern Russians.{{sfn|Balanovsky et al.|2008}} The highest frequency of R1a1a in Europe is observed in Sorbs (63%),{{sfn|Behar et al.|2003}} a West Slavic ethnic group, followed by Hungarians (60%).{{sfn|Semino et al.|2000}} Other groups with significant R1a1a, ranging from 27% to up to 58%, include Czechs, Poles, Slovenians, Slovaks, Moldovans, Belarusians, Rusyns, Ukrainians, and Russians.{{sfn|Balanovsky et al.|2008}}{{sfn|Behar et al.|2003}}{{sfn|Semino et al.|2000}} R1a frequency decreases in northeastern Russian populations down to 20%–30%, in contrast to central-southern Russia, where its frequency is twice as high. In the Baltics, R1a1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).{{sfn|Kasperaviciūte |Kucinskas |Stoneking |2005}}{{sfn|Battaglia et al.|2008}}{{sfn|Rosser et al.|2000}}{{sfn|Semino et al.|2000}}{{sfn|Tambets et al.|2004}}

There is also a significant presence in peoples of Germanic descent, with highest levels in Norway, Sweden and Iceland, where between 20 and 30% of men are in R1a1a.{{sfn|Bowden et al.|2008}}{{sfn|Dupuy et al.|2005}} Vikings and Normans may have also carried the R1a1a lineage further out, accounting for at least part of the small presence in the British Isles, the Canary Islands, and Sicily.{{sfn|Passarino et al.|2002}}{{sfn|Capelli et al.|2003}} Haplogroup R1a1a averages between 10 and 30% in Germans, with a peak in Rostock at 31.3%.{{sfn|Kayser et al.|2005}} R1a1a is found at a very low frequency among Dutch people (3.7%){{sfn|Semino et al.|2000}} and is virtually absent in Danes.{{Cite journal |last1=Sanchez |first1=J |last2=Børsting |first2=C |last3=Hallenberg |first3=C |last4=Buchard |first4=A |last5=Hernandez |first5=A |last6=Morling |first6=N |year=2003 |title=Multiplex PCR and minisequencing of SNPs—a model with 35 Y chromosome SNPs |journal=Forensic Science International |volume=137 |issue=1 |pages=74–84 |doi=10.1016/S0379-0738(03)00299-8 |pmid=14550618 |ref={{harvid|Sanchez et al.|2003}}}}

In Southern Europe R1a1a is not common, but significant levels have been found in pockets, such as in the Pas Valley in Northern Spain, areas of Venice, and Calabria in Italy.{{sfn|Scozzari et al.|2001}}{{Better source needed|date=January 2021}} The Balkans shows wide variation between areas with significant levels of R1a1a, for example 36–39% in Slovenia,{{cite journal|last1=Underhill|first1=Peter A.|title=The phylogenetic and geographic structure of Y-chromosome haplogroup R1a|journal=European Journal of Human Genetics|date=January 1, 2015|volume=23|issue=1|pages=124–131|doi=10.1038/ejhg.2014.50 |language=en|pmid=24667786|pmc=4266736}} 27–34% in Croatia,{{sfn|Battaglia et al.|2008}}{{cite journal|author=L. Barać|title=Y chromosomal heritage of Croatian population and its island isolates|journal=European Journal of Human Genetics|volume=11|issue=7|date=2003|pmid=12825075|doi=10.1038/sj.ejhg.5200992|ref={{harvid|Barać et al.|2003}}|display-authors=etal|pages=535–42|s2cid=15822710|doi-access=free}}{{cite journal|author=S. Rootsi|title=Phylogeography of Y-Chromosome Haplogroup I Reveals Distinct Domains of Prehistoric Gene Flow in Europe|url=http://evolutsioon.ut.ee/publications/Rootsi2004.pdf|journal=American Journal of Human Genetics|volume=75|issue=1|date=2004|pmid=15162323|pmc=1181996|doi=10.1086/422196|ref={{harvid|Rootsi et al.|2004}}|display-authors=etal|pages=128–137|access-date=February 13, 2021|archive-date=September 5, 2020|archive-url=https://web.archive.org/web/20200905162020/http://evolutsioon.ut.ee/publications/Rootsi2004.pdf|url-status=dead}}{{cite journal|author=M. Peričić|title=High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations|journal=Molecular Biology and Evolution|volume=22|issue=10|pages=1964–75|date=2005|pmid=15944443|doi=10.1093/molbev/msi185|ref={{harvid|Peričić et al.|2005}}|display-authors=etal|doi-access=free}}{{cite journal|author=M. Peričić|title=Review of Croatian Genetic Heritage as Revealed by Mitochondrial DNA and Y Chromosomal Lineages|journal=Croatian Medical Journal|volume=46|issue=4|date=2005|ref={{harvid|Peričić(2) et al.|2005}}|display-authors=etal|pmid=16100752|pages=502–513}} and over 30% in Greek Macedonia, but less than 10% in Albania, Kosovo and parts of Greece south of Olympus gorge.{{sfn|Pericić et al.|2005}}{{sfn|Rosser et al.|2000}}{{sfn|Semino et al.|2000}}

R1a is virtually composed only of the Z284 subclade in Scandinavia. In Slovenia, the main subclade is Z282 (Z280 and M458), although the Z284 subclade was found in one sample of a Slovenian. There is a negligible representation of Z93 in Turkey, 12,1%{{sfn|Underhill et al.|2014}} West Slavs and Hungarians are characterized by a high frequency of the subclade M458 and a low Z92, a subclade of Z280. Hundreds of Slovenian samples and Czechs lack the Z92 subclade of Z280, while Poles, Slovaks, Croats and Hungarians only show a very low frequency of Z92.{{sfn|Underhill et al.|2014}} The Balts, East Slavs, Serbs, Macedonians, Bulgarians and Romanians demonstrate a ratio Z280>M458 and a high, up to a prevailing share of Z92.{{sfn|Underhill et al.|2014}} Balts and East Slavs have the same subclades and similar frequencies in a more detailed phylogeny of the subclades.{{cite web |title=Untitled |website=pereformat.ru |language=ru |url=http://pereformat.ru/wp-content/uploads/2015/02/russian-plain-01.jpg |access-date=May 29, 2017 |archive-date=March 15, 2016 |archive-url=https://web.archive.org/web/20160315183214/http://pereformat.ru/wp-content/uploads/2015/02/russian-plain-01.jpg |url-status=live }}{{cite web |title=Untitled |website=www.rodstvo.ru |url=http://www.rodstvo.ru/forum/index.php?act=attach&type=post&id=1299 |access-date=May 29, 2017 |archive-date=September 16, 2021 |archive-url=https://web.archive.org/web/20210916214250/https://www.rodstvo.ru/forum/index.php?act=attach&type=post&id=1299 |url-status=dead }} The Russian geneticist Oleg Balanovsky speculated that there is a predominance of the assimilated pre-Slavic substrate in the genetics of East and West Slavic populations, according to him the common genetic structure which contrasts East Slavs and Balts from other populations may suggest the explanation that the pre-Slavic substrate of the East and West Slavs consisted most significantly of Baltic-speakers, which at one point predated the Slavs in the cultures of the Eurasian steppe according to archaeological and toponymic references.{{refn|group=note|{{harvp|Балановский|2015|p=[https://books.google.com/books?id=sNYPCwAAQBAJ&pg=PA208 208]}} {{lang in|ru}} {{lang|ru|Прежде всего, это преобладание в славянских популяциях дославянского субстрата — двух ассимилированных ими генетических компонентов – восточноевропейского для западных и восточных славян и южноевропейского для южных славян...Можно с осторожностью предположить, что ассимилированный субстратмог быть представлен по преимуществу балтоязычными популяциями. Действительно, археологические данные указыва ют на очень широкое распространение балтских групп перед началом расселения славян. Балтскийсубстрату славян (правда, наряду с финно-угорским) выявляли и антропологи. Полученные нами генетические данные — и на графиках генетических взаимоотношений, и по доле общих фрагментов генома — указывают, что современные балтские народы являются ближайшими генетически ми соседями восточных славян. При этом балты являются и лингвистически ближайшими род ственниками славян. И можно полагать, что к моменту ассимиляции их генофонд не так сильно отличался от генофонда начавших свое широкое расселение славян. Поэтому если предположить,что расселяющиеся на восток славяне ассимилировали по преимуществу балтов, это может объяснить и сходство современных славянских и балтских народов друг с другом, и их отличия от окружающих их не балто-славянских групп Европы...В работе высказывается осторожное предположение, что ассимилированный субстрат мог быть представлен по преимуществу балтоязычными популяциями. Действительно, археологические данные указывают на очень широкое распространение балтских групп перед началом расселения славян. Балтский субстрат у славян (правда, наряду с финно-угорским) выявляли и антропологи. Полученные в этой работе генетические данные — и на графиках генетических взаимоотношений, и по доле общих фрагментов генома — указывают, что современные балтские народы являются ближайшими генетическими соседями восточных славян.}}}}

{{harvp|Zerjal et al.|2002}} found R1a1a in 64% of a sample of the Tajiks of Tajikistan and 63% of a sample of the Kyrgyz of Kyrgyzstan.{{sfn|Zerjal et al.|2002}}

{{harvp|Haber et al.|2012}} found R1a1a-M17 in 26.0% (53/204) of a set of samples from Afghanistan, including 60% (3/5) of a sample of Nuristanis, 51.0% (25/49) of a sample of Pashtuns, 30.4% (17/56) of a sample of Tajiks, 17.6% (3/17) of a sample of Uzbeks, 6.7% (4/60) of a sample of Hazaras, and in the only sampled Turkmen individual.{{sfn|Haber et al.|2012}}

{{harvp|Di Cristofaro et al.|2013}} found R1a1a-M198/M17 in 56.3% (49/87) of a pair of samples of Pashtuns from Afghanistan (including 20/34 or 58.8% of a sample of Pashtuns from Baghlan and 29/53 or 54.7% of a sample of Pashtuns from Kunduz), 29.1% (37/127) of a pool of samples of Uzbeks from Afghanistan (including 28/94 or 29.8% of a sample of Uzbeks from Jawzjan, 8/28 or 28.6% of a sample of Uzbeks from Sar-e Pol, and 1/5 or 20% of a sample of Uzbeks from Balkh), 27.5% (39/142) of a pool of samples of Tajiks from Afghanistan (including 22/54 or 40.7% of a sample of Tajiks from Balkh, 9/35 or 25.7% of a sample of Tajiks from Takhar, 4/16 or 25.0% of a sample of Tajiks from Samangan, and 4/37 or 10.8% of a sample of Tajiks from Badakhshan), 16.2% (12/74) of a sample of Turkmens from Jawzjan, and 9.1% (7/77) of a pair of samples of Hazara from Afghanistan (including 7/69 or 10.1% of a sample of Hazara from Bamiyan and 0/8 or 0% of a sample of Hazara from Balkh).{{sfn|Di Cristofaro et al.|2013}}

{{harvp|Malyarchuk et al.|2013}} found R1a1-SRY10831.2 in 30.0% (12/40) of a sample of Tajiks from Tajikistan.{{sfn|Malyarchuk et al.|2013}}

{{harvp|Ashirbekov et al.|2017}} found R1a-M198 in 6.03% (78/1294) of a set of samples of Kazakhs from Kazakhstan. R1a-M198 was observed with greater than average frequency in the study's samples of the following Kazakh tribes: 13/41 = 31.7% of a sample of Suan, 8/29 = 27.6% of a sample of Oshaqty, 6/30 = 20.0% of a sample of Qozha, 4/29 = 13.8% of a sample of Qypshaq, 1/8 = 12.5% of a sample of Tore, 9/86 = 10.5% of a sample of Jetyru, 4/50 = 8.0% of a sample of Argyn, 1/13 = 7.7% of a sample of Shanyshqyly, 8/122 = 6.6% of a sample of Alimuly, 3/46 = 6.5% of a sample of Alban. R1a-M198 also was observed in 5/42 = 11.9% of a sample of Kazakhs of unreported tribal affiliation.{{sfn|Ashirbekov et al.|2017}}

In South Asia, R1a1a has often been observed in a number of demographic groups.{{sfn|Sahoo et al.|2006}}{{sfn|Sengupta|2006}}

In India, high frequencies of this haplogroup is observed in West Bengal Brahmins (72%) in the east,{{sfn|Sengupta|2006}} Bhanushali (67%) and Gujarat Lohanas (60%) in the west,{{sfn|Underhill et al.|2009}} Uttar Pradesh Brahmins (68%), Punjab/Haryana Khatris (67%) and Ahirs (63%) in the north,{{sfn|Sharma et al.|2009}}{{sfn|Sengupta|2006}}{{sfn|Underhill et al.|2009}} and Karnataka Medars (39%) in the south.{{sfn|Shah|2011}} It has also been found in several South Indian Dravidian-speaking Adivasis including the Chenchu (26%) of Andhra Pradesh and Kota of Andhra Pradesh (22.58%){{sfn|Arunkumar|2012}} and the Kallar of Tamil Nadu suggesting that R1a1a is widespread in Tribal Southern Indians.{{sfn|Kivisild et al.|2003}}

Besides these, studies show high percentages in regionally diverse groups such as Manipuris (50%){{sfn|Underhill et al.|2009}} to the extreme North East and among Punjabis (47%){{sfn|Kivisild et al.|2003}} to the extreme North West.

In Pakistan it is found at 80% among Yusufzai tribe of Pashtuns (51%) from Swat District,{{cite journal |title=Contrasting maternal and paternal genetic histories among five ethnic groups from Khyber Pakhtunkhwa, Pakistan |author=Tariq, Muhammad; Ahmad, Habib; Hemphill, Brian E.; Farooq, Umar; Schurr, Theodore G. |journal=Scientific Reports |volume=12 |issue=1 |pages=1027 |year=2022 |doi=10.1038/s41598-022-05076-3 |pmid=35046511 |pmc=8770644 |bibcode=2022NatSR..12.1027T }} 71% among the Mohanna community in Sindh province to the south and 46% among the Baltis of Gilgit-Baltistan to the north.{{sfn|Underhill et al.|2009}}

Among the Sinhalese of Sri Lanka, 23% were found to be R1a1a (R-SRY1532) positive.{{Cite book |chapter-url=http://evolutsioon.ut.ee/publications/Kivisild2003a.pdf |author=Toomas Kivisild |author2=Siiri Rootsi |author3=Mait Metspalu |author4=Ene Metspalu |author5=Juri Parik |author6=Katrin Kaldma |author7=Esien Usanga |author8=Sarabjit Mastana |author9=Surinder S. Papiha |author10=Richard Villems |chapter=The Genetics of Language and Farming Spread in India |access-date=December 20, 2019 |title=Examining the farming/language dispersal hypothesis |editor=P. Bellwwood |editor2=C. Renfrew |series=McDonald Institute Monographs |publisher=Cambridge University |pages=215–222 |archive-date=February 19, 2006 |archive-url=https://web.archive.org/web/20060219054915/http://evolutsioon.ut.ee/publications/Kivisild2003a.pdf |url-status=dead }} Hindus of Chitwan District in the Terai region Nepal show it at 69%.{{sfn|Fornarino et al.|2009}}

The frequency of R1a1a is comparatively low among some Turkic-speaking groups like Yakuts, yet levels are higher (19 to 28%) in certain Turkic or Mongolic-speaking groups of Northwestern China, such as the Bonan, Dongxiang, Salar, and Uyghurs.{{sfn|Wells|2001}}{{sfn|Wang et al.|2003}}{{sfn|Zhou et al.|2007}}

A Chinese paper published in 2018 found R1a-Z94 in 38.5% (15/39) of a sample of Keriyalik Uyghurs from Darya Boyi / Darya Boye Village, Yutian County, Xinjiang (于田县达里雅布依乡), R1a-Z93 in 28.9% (22/76) of a sample of Dolan Uyghurs from Horiqol township, Awat County, Xinjiang (阿瓦提县乌鲁却勒镇), and R1a-Z93 in 6.3% (4/64) of a sample of Loplik Uyghurs from Karquga / Qarchugha Village, Yuli County, Xinjiang (尉犁县喀尔曲尕乡). R1a(xZ93) was observed only in one of 76 Dolan Uyghurs.{{sfn|Liu Shu-hu et al.|2018}} Note that Darya Boyi Village is located in a remote oasis formed by the Keriya River in the Taklamakan Desert. A 2011 Y-DNA study found Y-dna R1a1 in 10% of a sample of southern Hui people from Yunnan, 1.6% of a sample of Tibetan people from Tibet (Tibet Autonomous Region), 1.6% of a sample of Xibe people from Xinjiang, 3.2% of a sample of northern Hui from Ningxia, 9.4% of a sample of Hazak (Kazakhs) from Xinjiang, and rates of 24.0%, 22.2%, 35.2%, 29.2% in 4 different samples of Uyghurs from Xinjiang, 9.1% in a sample of Mongols from Inner Mongolia. A different subclade of R1 was also found in 1.5% of a sample of northern Hui from Ningxia.{{sfn|Zhong et al.|2011}} in the same study there were no cases of R1a detected at all in 6 samples of Han Chinese in Yunnan, 1 sample of Han in Guangxi, 5 samples of Han in Guizhou, 2 samples of Han in Guangdong, 2 samples of Han in Fujian, 2 samples of Han in Zhejiang, 1 sample of Han in Shanghai, 1 samples of Han in Jiangxi, 2 samples of Han in Hunan, 1 sample of Han in Hubei, 2 samples of Han in Sichuan, 1 sample of Han in Chongqing, 3 samples of Han in Shandong, 5 samples of Han in Gansu, 3 samples of Han in Jilin and 2 samples of Han in Heilongjiang.{{cite journal |last1=Zhong |first1=Hua |last2=Shi |first2= Hong |last3=Qi |first3=Xue-Bin |last4=Duan |first4=Zi-Yuan |last5=Tan |first5= Ping-Ping |last6= Jin|first6=Li |last7=Su |first7=Bing |last8=Ma |first8=Runlin Z. |date=2011 |title= Extended Y Chromosome Investigation Suggests Postglacial Migrations of Modern Humans into East Asia via the Northern Route|journal=Molecular Biology and Evolution |volume=28 |issue=1 |pages=717–727 |doi=10.1093/molbev/msq247 |doi-access=free |pmid=20837606}} 40% of Salars, 45.2% of Tajiks of Xinjiang, 54.3% of Dongxiang, 60.6% of Tatars and 68.9% of Kyrgyz in Xinjiang in northwestern China tested in one sample had R1a1-M17. Bao'an (Bonan) had the most haplogroup diversity of 0.8946±0.0305 while the other ethnic minorities in northwestern China had a high haplogroup diversity like Central Asians, of 0.7602±0.0546.{{cite journal |last1=Shou |first1=Wei-Hua |last2= Qiao|first2= Wn-Fa |last3=Wei |first3=Chuan-Yu |last4= Dong|first4= Yong-Li |last5=Tan |first5=Si-Jie |last6=Shi |first6=Hong |last7=Tang |first7=Wen-Ru |last8= Xiao |first8=Chun-Jie |date=2010|title=Y-chromosome distributions among populations in Northwest China identify significant contribution from Central Asian pastoralists and lesser influence of western Eurasians |journal= Journal of Human Genetics|volume=55 |issue= 5|pages= 314–322 |doi=10.1038/jhg.2010.30 |pmid=20414255 |s2cid=23002493 |doi-access=free }}

In Eastern Siberia, R1a1a is found among certain indigenous ethnic groups including Kamchatkans and Chukotkans, and peaking in Itel'man at 22%.{{sfn|Lell et al.|2002}}

Y-haplogroups R1a-M420 and R2-M479 are found in Ede (8.3% and 4.2%) and Giarai (3.7% and 3.7%) peoples in Vietnam. The Cham additionally have haplogroups R-M17 (13.6%) and R-M124 (3.4%).

R1a1a1b2a2a (R-Z2123) and R1a1 are found in Khmer peoples from Thailand (3.4%) and Cambodia (7.2%) respectively. Haplogroup R1a1a1b2a1b (R-Y6) is also found among Kuy peoples (5%).

According to Changmai et al. (2022), these haplogroup frequencies originate from South Asians, who left a cultural and genetic legacy in Southeast Asia since the first millennium CE.{{Cite journal |last1=Changmai |first1=Piya |last2=Jaisamut |first2=Kitipong |last3=Kampuansai |first3=Jatupol |last4=Kutanan |first4=Wibhu |display-authors=3 |date=2022 |title=Indian genetic heritage in Southeast Asian populations |journal=PLOS Genetics |volume=18 |issue=2 |pages=e1010036 |doi=10.1371/journal.pgen.1010036 |pmc=8853555 |pmid=35176016 |doi-access=free}}

R1a1a has been found in various forms, in most parts of Western Asia, in widely varying concentrations, from almost no presence in areas such as Jordan, to much higher levels in parts of Kuwait and Iran. The Shimar (Shammar) Bedouin tribe in Kuwait show the highest frequency in the Middle East at 43%.{{sfn|Mohammad et al.|2009}}{{sfn|Nasidze et al.|2004}}{{sfn|Nasidze et al.|2005}}

{{Harvnb|Wells|2001}}, noted that in the western part of the country, Iranians show low R1a1a levels, while males of eastern parts of Iran carried up to 35% R1a1a. {{Harvnb|Nasidze et al.|2004}} found R1a1a in approximately 20% of Iranian males from the cities of Tehran and Isfahan. {{Harvnb|Regueiro|2006}} in a study of Iran, noted much higher frequencies in the south than the north.

A newer study has found 20.3% R-M17* among Kurdish samples which were taken in the Kurdistan Province in western Iran, 19% among Azerbaijanis in West Azerbaijan, 9.7% among Mazandaranis in North Iran in the province of Mazandaran, 9.4% among Gilaks in province of Gilan, 12.8% among Persian and 17.6% among Zoroastrians in Yazd, 18.2% among Persians in Isfahan, 20.3% among Persians in Khorasan, 16.7% Afro-Iranians, 18.4% Qeshmi "Gheshmi", 21.4% among Persian Bandari people in Hormozgan and 25% among the Baloch people in Sistan and Baluchestan Province.{{sfn|Grugni et al.|2012}}

{{harvp|Di Cristofaro et al.|2013}} found haplogroup R1a in 9.68% (18/186) of a set of samples from Iran, though with a large variance ranging from 0% (0/18) in a sample of Iranians from Tehran to 25% (5/20) in a sample of Iranians from Khorasan and 27% (3/11) in a sample of Iranians of unknown provenance. All Iranian R1a individuals carried the M198 and M17 mutations except one individual in a sample of Iranians from Gilan (n=27), who was reported to belong to R1a-SRY1532.2(xM198, M17).{{sfn|Di Cristofaro et al.|2013}}

{{harvp|Malyarchuk et al.|2013}} found R1a1-SRY10831.2 in 20.8% (16/77) of a sample of Persians collected in the provinces of Khorasan and Kerman in eastern Iran, but they did not find any member of this haplogroup in a sample of 25 Kurds collected in the province of Kermanshah in western Iran.{{sfn|Malyarchuk et al.|2013}}

Further to the north of these Western Asian regions on the other hand, R1a1a levels start to increase in the Caucasus, once again in an uneven way. Several populations studied have shown no sign of R1a1a, while highest levels so far discovered in the region appears to belong to speakers of the Karachay-Balkar language among whom about one quarter of men tested so far are in haplogroup R1a1a.{{sfn|Underhill et al.|2009}}

{{further|Conversion table for Y chromosome haplogroups}}

The historic naming system commonly used for R1a was inconsistent in different published sources, because it changed often; this requires some explanation.

In 2002, the Y Chromosome Consortium (YCC) proposed a new naming system for haplogroups {{harv|YCC|2002}}, which has now become standard. In this system, names with the format "R1" and "R1a" are "phylogenetic" names, aimed at marking positions in a family tree. Names of SNP mutations can also be used to name clades or haplogroups. For example, as M173 is currently the defining mutation of R1, R1 is also R-M173, a "mutational" clade name. When a new branching in a tree is discovered, some phylogenetic names will change, but by definition all mutational names will remain the same.

The widely occurring haplogroup defined by mutation M17 was known by various names, such as "Eu19", as used in {{harv|Semino et al.|2000}} in the older naming systems. The 2002 YCC proposal assigned the name R1a to the haplogroup defined by mutation SRY1532.2. This included Eu19 (i.e. R-M17) as a subclade, so Eu19 was named R1a1. Note, SRY1532.2 is also known as SRY10831.2{{citation needed|date=December 2012}} The discovery of M420 in 2009 has caused a reassignment of these phylogenetic names.({{harvnb|Underhill et al.|2009}} and {{harvnb|ISOGG|2012}}) R1a is now defined by the M420 mutation: in this updated tree, the subclade defined by SRY1532.2 has moved from R1a to R1a1, and Eu19 (R-M17) from R1a1 to R1a1a.

More recent updates recorded at the ISOGG reference webpage involve branches of R-M17, including one major branch, R-M417.

{{clear}}

{{Clear right}}

{{columns-list|colwidth=22em|

• List of R1a frequency by population
• Genetic history of Europe
• Genetics and archaeogenetics of South Asia
• Haplogroup Q-M242 (Y-DNA)
• Human Y-chromosome DNA haplogroups
• Neanderthal Y-chromosome DNA haplogroups
• Y-chromosome haplogroups in populations of the world

}}

{{columns-list|colwidth=18em|

• R-L21
• R-L295
• R-M124
• R-M167
• R-M17
• R-M173
• R-M207
• R-M342
• R-M420
• R-M479
• R-U106

}}

{{Y-DNA}}

{{reflist|group=note}}

{{reflist|32em}}

{{refbegin|40em}}

• {{Cite journal |ref={{sfnref|Allentoft et al.|2015}} |last1=Allentoft |first1=Morten E. |last2=Sikora |first2=Martin |last3=Sjögren |first3=Karl-Göran |last4=Rasmussen |first4=Simon |last5=Rasmussen |first5=Morten |last6=Stenderup |first6=Jesper |last7=Damgaard |first7=Peter B. |last8=Schroeder |first8=Hannes |last9=Ahlström |first9=Torbjörn |last10=Vinner |first10=Lasse |last11=Malaspinas |first11=Anna-Sapfo |last12=Margaryan |first12=Ashot |last13=Higham |first13=Tom |last14=Chivall |first14=David |last15=Lynnerup |first15=Niels |last16=Harvig |first16=Lise |last17=Baron |first17=Justyna |last18=Casa |first18=Philippe Della |last19=Dąbrowski |first19=Paweł |last20=Duffy |first20=Paul R. |last21=Ebel |first21=Alexander V. |last22=Epimakhov |first22=Andrey |last23=Frei |first23=Karin |last24=Furmanek |first24=Mirosław |last25=Gralak |first25=Tomasz |last26=Gromov |first26=Andrey |last27=Gronkiewicz |first27=Stanisław |last28=Grupe |first28=Gisela |last29=Hajdu |first29=Tamás |last30=Jarysz |first30=Radosław |s2cid=4399103 |display-authors=8 |title=Population genomics of Bronze Age Eurasia |journal=Nature |volume=522 |issue=7555 |pages=167–172 |year=2015 |url=https://depot.ceon.pl/handle/123456789/13155 |bibcode=2015Natur.522..167A |doi=10.1038/nature14507 |pmid=26062507 |access-date=July 23, 2019 |archive-date=July 13, 2019 |archive-url=https://web.archive.org/web/20190713182031/https://depot.ceon.pl/handle/123456789/13155 |url-status=live }}
• {{Citation |last=Anthony |first=David W. |year=2007 |title=The Horse The Wheel And Language. How Bronze-Age Riders From the Eurasian Steppes Shaped The Modern World |publisher=Princeton University Press }}
• {{cite journal |last1=Anthony |first1=David |author-link1=David W. Anthony |date=Spring–Summer 2019 |title=Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard |url=https://www.academia.edu/39985565 |access-date=January 9, 2020 |journal=Journal of Indo-European Studies |volume=47 |issue=1–2 |archive-date=May 3, 2024 |archive-url=https://web.archive.org/web/20240503032509/https://www.academia.edu/39985565/Archaeology_Genetics_and_Language_in_the_Steppes_A_Comment_on_Bomhard |url-status=live }}
• {{Citation |last1=Anthony |first1=David |last2=Ringe |first2=Don |year=2015 |title=The Indo-European Homeland from Linguistic and Archaeological Perspectives |journal=Annual Review of Linguistics |doi=10.1146/annurev-linguist-030514-124812 |volume=1 |pages=199–219|doi-access=free }}
• {{cite journal| last1=Shah| year=2011| doi=10.1016/j.ajhg.2011.05.030| title=Indian Siddis: African Descendants with Indian Admixture| first1=A. M.| last2=Tamang| first2=R.| last3=Moorjani| first3=P.| last4=Rani| first4=D. S.| last5=Govindaraj| first5=P.| last6=Kulkarni| first6=G.| last7=Bhattacharya| first7=T.| last8=Mustak| first8=M. S.| last9=Bhaskar| first9=L. V. K. S.| last10=Reddy| first10=A. G.| last11=Gadhvi| first11=D.| last12=Gai| first12=P. B.| last13=Chaubey| first13=G.| last14=Patterson| first14=N.| last15=Reich| first15=D.| last16=Tyler-Smith| first16=C.| last17=Singh| first17=L.| last18=Thangaraj| first18=K.| journal=The American Journal of Human Genetics| volume=89| issue=1| pages=154–61| ref={{harvid|Shah|2011}}| pmid=21741027| pmc=3135801}}
• {{Cite journal |last1=ArunKumar |first1=G |last2=Soria-Hernanz | first2=DF |last3=Kavitha |first3=VJ |last4=Arun |first4=VS |last5=Syama |first5=A |last6=Ashokan |first6=KS |title=Population Differentiation of Southern Indian Male Lineages Correlates with Agricultural Expansions Predating the Caste System |year=2012 |journal= PLOS ONE |volume=7 |issue=11 |page=e50269|doi=10.1371/journal.pone.0050269 |pmid=23209694 |pmc=3508930 |ref={{harvid |Arunkumar |2012}}|bibcode=2012PLoSO...750269A |doi-access=free }}
• {{cite journal |ref={{sfnref|Ashirbekov et al.|2017}} |first=E. E. |last=Ashirbekov |display-authors=et al |date=2017 |title=Distribution of Y-Chromosome Haplogroups of the Kazakh from the South Kazakhstan, Zhambyl, and Almaty Regions |journal=Reports of the National Academy of Sciences of the Republic of Kazakhstan |volume=6 |issue=316 |pages=85–95 |url=http://nblib.library.kz/elib/library.kz/jurnal/%D0%94%D0%BE%D0%BA%D0%BB%D0%B0%D0%B4_06_2017%20(2)/12-%20Biology%20E.E.Ashirbekov0617.pdf |access-date=June 4, 2020 |archive-date=September 23, 2021 |archive-url=https://web.archive.org/web/20210923024313/http://nblib.library.kz/elib/library.kz/jurnal/%D0%94%D0%BE%D0%BA%D0%BB%D0%B0%D0%B4_06_2017%20(2)/12-%20Biology%20E.E.Ashirbekov0617.pdf |url-status=dead }}
• {{Cite journal |vauthors=Balanovsky O, Rootsi S, Pshenichnov A, Kivisild T, Churnosov M, Evseeva I, Pocheshkhova E, Boldyreva M, Yankovsky N, Balanovska E, Villems R |title=Two Sources of the Russian Patrilineal Heritage in Their Eurasian Context |journal=American Journal of Human Genetics |volume=82 |issue=1 |pages=236–250 |year=2008 |doi=10.1016/j.ajhg.2007.09.019 |pmc=2253976 |pmid=18179905 |display-authors=8 |ref={{sfnref|Balanovsky et al.|2008}} }}
• {{Cite book|last1=Балановский|first1=О. П.|title=Генофонд Европы|url=https://books.google.com/books?id=sNYPCwAAQBAJ|publisher=KMK Scientific Press|language=ru|isbn=9785990715707|date=November 30, 2015|access-date=August 8, 2020|archive-date=May 3, 2024|archive-url=https://web.archive.org/web/20240503032542/https://books.google.com/books?id=sNYPCwAAQBAJ|url-status=live}}
• {{Cite journal |vauthors=Battaglia V, Fornarino S, Al-Zahery N, Olivieri A, Pala M, Myres NM, King RJ, Rootsi S, Marjanovic D, Primorac D, Hadziselimovic R, Vidovic S, Drobnic K, Durmishi N, Torroni A, Santachiara-Benerecetti AS, Underhill PA, Semino O |year=2008 |title=Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe |volume=17 |issue=6 |pages=820–30 |journal=European Journal of Human Genetics |doi=10.1038/ejhg.2008.249 |pmc=2947100 |pmid=19107149 |display-authors=8 |ref={{sfnref|Battaglia et al.|2008}} }}
• {{Cite journal |vauthors=Behar D, Thomas MG, Skorecki K, Hammer MF, Bulygina E, Rosengarten D, Jones AL, Held K, Moses V, Goldstein D, Bradman N, Weale ME |display-authors=8 |year=2003 |title=Multiple Origins of Ashkenazi Levites: Y Chromosome Evidence for Both Near Eastern and European Ancestries |journal=American Journal of Human Genetics |volume=73 |issue=4 |pages=768–779 |pmid=13680527 |pmc=1180600 |doi=10.1086/378506 |doi-access=free |ref={{sfnref|Behar et al.|2003}} |url=http://www.ucl.ac.uk/tcga/tcgapdf/Behar-AJHG-03.pdf |access-date=April 23, 2007 |archive-date=April 17, 2007 |archive-url=https://web.archive.org/web/20070417221913/http://www.ucl.ac.uk/tcga/tcgapdf/Behar-AJHG-03.pdf |url-status=live }}
• {{Cite journal |vauthors=Bowden GR, Balaresque P, King TE, Hansen Z, Lee AC, Pergl-Wilson G, Hurley E, Roberts SJ, Waite P, Jesch J, Jones AL, Thomas MG, Harding SE, Jobling MA |title=Excavating Past Population Structures by Surname-Based Sampling: The Genetic Legacy of the Vikings in Northwest England |journal=Molecular Biology and Evolution |year=2008 |volume=25 |pmid=18032405 |issue=2 |pmc=2628767 |pages=301–309 |doi=10.1093/molbev/msm255 |display-authors=8 |ref={{sfnref|Bowden et al.|2008}} }}
• {{cite journal |ref={{sfnref|Brandit et al.|2013}} |last1=Brandit |first1=G. |collaboration=The Genographic Consortium |year=2013 |title=Ancient DNA Reveals Key Stages in the Formation of Central European Mitochondrial Genetic Diversity |journal=Science |volume=342 |issue=6155 |pages=257–261 |doi= 10.1126/science.1241844 |pmid= 24115443 |pmc= 4039305 |bibcode= 2013Sci...342..257B }}
• {{Cite journal |vauthors=Capelli C, Redhead N, Abernethy JK, Gratrix F, Wilson JF, Moen T, Hervig T, Richards M, Stumpf MP, Underhill PA, Bradshaw P, Shaha A, Thomas MG, Bradman N, Goldstein DB |s2cid=526263 |title=A Y Chromosome Census of the British Isles |journal=Current Biology |volume=13 |issue=11 |pages=979–84 |year=2003 |pmid=12781138 |doi=10.1016/S0960-9822(03)00373-7 |bibcode=2003CBio...13..979C |url=https://www.pure.ed.ac.uk/ws/files/10690515/A_Y_chromosome_census_of_the_British_Isles.pdf |display-authors=8 |ref={{sfnref|Capelli et al.|2003}} |access-date=August 8, 2020 |archive-date=July 8, 2021 |archive-url=https://web.archive.org/web/20210708152834/https://www.pure.ed.ac.uk/ws/files/10690515/A_Y_chromosome_census_of_the_British_Isles.pdf |url-status=live }} also at {{cite web |url=http://www.ucl.ac.uk/tcga/tcgapdf/capelli-CB-03.pdf |title=University College London |access-date=January 29, 2007 |archive-date=January 17, 2017 |archive-url=https://web.archive.org/web/20170117022551/http://www.ucl.ac.uk/tcga/tcgapdf/capelli-CB-03.pdf |url-status=live }}
• {{cite journal |ref={{sfnref|Chunxiang Li et al.|2010}} |author=Chunxiang Li |author2=Hongjie Li |author3=Yinqiu Cui |author4=Chengzhi Xie |author5=Dawei Cai |author6=Wenying Li |author7=Victor H Mair |author8=Zhi Xu |author9=Quanchao Zhang |author10=Idelisi Abuduresule |author11=Li Jin |author12=Hong Zhu |author13=Hui Zhou |display-authors=8 |title=Evidence that a West-East admixed population lived in the Tarim Basin as early as the early Bronze Age |journal=BMC Biology |volume=8 |issue=1 |year=2010 |pages=15 |doi=10.1186/1741-7007-8-15 |pmid=20163704 |pmc=2838831 |doi-access=free }}
• {{cite journal |ref={{sfnref| Di Cristofaro et al.|2013}} |vauthors= Di Cristofaro J, Pennarun E, Mazières S, Myres NM, Lin AA, Temori SA, Metspalu M, Metspalu E, Witzel M, King RJ, Underhill PA, Villems R, Chiaroni J |display-authors= 8 |year=2013 |title= Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge |journal= PLOS ONE |volume=8 |issue=10 |at=e76748 |doi= 10.1371/journal.pone.0076748 |pmid= 24204668 |pmc= 3799995 |bibcode= 2013PLoSO...876748D|doi-access= free }}
• {{Cite journal |vauthors=Dupuy BM, Stenersen M, Lu TT, Olaisen B |year=2005 |title=Geographical heterogeneity of Y-chromosomal lineages in Norway |journal=Forensic Science International |volume=164 |issue=1 |pages=10–19 |pmid=16337760 |doi=10.1016/j.forsciint.2005.11.009 |url=http://www.freewebs.com/rus_anthro/Dupuy_2005_Geographical_heterogeneity_of_Y_chromosomal_lineages_in_Norway_FSI.pdf |ref={{sfnref|Dupuy et al.|2005}} |access-date=November 18, 2009 |archive-date=July 22, 2012 |archive-url=https://web.archive.org/web/20120722183015/http://www.freewebs.com/rus_anthro/Dupuy_2005_Geographical_heterogeneity_of_Y_chromosomal_lineages_in_Norway_FSI.pdf |url-status=live }}
• {{Cite journal |last1=Fornarino |first1=Simona |last2=Pala |first2=Maria |last3=Battaglia |first3=Vincenza |last4=Maranta |first4=Ramona |last5=Achilli |first5=Alessandro |last6=Modiano |first6=Guido |last7=Torroni |first7=Antonio |last8=Semino |first8=Ornella |last9=Santachiara-Benerecetti |first9=Silvana A. |display-authors=8 |year=2009 |title=Mitochondrial and Y-chromosome diversity of the Tharus (Nepal): a reservoir of genetic variation |journal=BMC Evolutionary Biology |volume=9 |issue=1 |page=154 |doi= 10.1186/1471-2148-9-154 |pmc= 2720951 |pmid= 19573232 |bibcode=2009BMCEE...9..154F |ref={{sfnref|Fornarino et al.|2009}} |doi-access=free }}
• {{cite journal |ref={{sfnref|Fu et al.|2016}} |last1=Fu |first1=Qiaomei |display-authors=etal |date=May 2, 2016 |title=The genetic history of Ice Age Europe |journal=Nature |volume=534 |issue=7606 |pages=200–205 |bibcode= 2016Natur.534..200F |doi= 10.1038/nature17993 |pmc=4943878 |pmid=27135931 |hdl=10211.3/198594}}
• {{cite journal |vauthors= Grugni V, Battaglia V, Kashani BH, Parolo S, Al-Zahery N, Achilli A, Olivieri A, Gandini F, Houshmand M, Sanati MH, Torroni A, Semino O |year=2012 |title=Ancient Migratory Events in the Middle East: New Clues from the Y-Chromosome Variation of Modern Iranians |journal=PLOS ONE |volume=7 |issue=7 |at=e41252 |doi= 10.1371/journal.pone.0041252 |doi-access=free |pmid= 22815981 |pmc= 3399854 |bibcode= 2012PLoSO...741252G |ref={{sfnref |Grugni et al.|2012}} }}
• {{Cite journal |author=Gwozdz |title=Y-STR Mountains in Haplospace, Part II: Application to Common Polish Clades |url=http://www.jogg.info/52/files/Gwozdz2.pdf |journal=Journal of Genetic Genealogy |volume=5 |issue=2 |year=2009 |access-date=November 28, 2009 |archive-date=July 21, 2011 |archive-url=https://web.archive.org/web/20110721115739/http://www.jogg.info/52/files/Gwozdz2.pdf |url-status=live }}
• {{Cite journal |ref={{sfnref|Haak et al.|2008}} |last1=Haak |first1=W. |last2=Brandt |first2=G. |last3=Jong |first3=H. N. d. |last4=Meyer |first4=C. |last5=Ganslmeier |first5=R. |last6=Heyd |first6=V. |last7=Hawkesworth |first7=C. |last8=Pike |first8=A. W. G. |last9=Meller |first9=H. |last10=Alt |first10=K. W. |title=Ancient DNA, Strontium isotopes, and osteological analyses shed light on social and kinship organization of the Later Stone Age |doi=10.1073/pnas.0807592105 |journal=Proceedings of the National Academy of Sciences |year=2008 |pmc=2587582 |volume=105 |issue=47 |pages=18226–18231 |pmid=19015520 |bibcode=2008PNAS..10518226H |display-authors=8|doi-access=free }}
• {{Cite journal |ref={{sfnref|Haak et al.|2015}} |last1= Haak |first1= Wolfgang |last2= Lazaridis |first2= Iosif |last3= Patterson |first3= Nick |last4= Rohland |first4= Nadin |last5= Mallick |first5= Swapan |last6= Llamas |first6= Bastien |last7= Brandt |first7= Guido |last8= Nordenfelt |first8= Susanne |last9= Harney |first9= Eadaoin |last10= Stewardson |first10= Kristin |last11= Fu |first11= Qiaomei |last12= Mittnik |first12= Alissa |last13= Bánffy |first13= Eszter |last14= Economou |first14= Christos |last15= Francken |first15= Michael |last16= Friederich |first16= Susanne |last17= Pena |first17= Rafael Garrido |last18= Hallgren |first18= Fredrik |last19= Khartanovich |first19= Valery |last20= Khokhlov |first20= Aleksandr |last21= Kunst |first21= Michael |last22= Kuznetsov |first22= Pavel |last23= Meller |first23= Harald |last24= Mochalov |first24= Oleg |last25= Moiseyev |first25= Vayacheslav |last26= Nicklisch |first26= Nicole |last27= Pichler |first27= Sandra L. |last28= Risch |first28= Roberto |last29= Rojo Guerra |first29= Manuel A. |last30= Roth |first30= Christina |first31=Anna |last31=Szécsényi-Nagy |first32=Joachim |last32=Wahl |first33=Matthias |last33=Meyer |first34=Johannes |last34=Krause |first35=Dorcas |last35=Brown |first36=David |last36=Anthony |first37=Alan |last37=Cooper |first38=Kurt Werner |last38=Alt |first39=David |last39=Reich |display-authors=8 |title= Massive migration from the steppe is a source for Indo-European languages in Europe |journal= Nature|volume=522 |issue=7555 |year=2015 |pages= 207–211 |arxiv= 1502.02783 |bibcode= 2015Natur.522..207H |pmid= 25731166 |pmc= 5048219 |biorxiv= 10.1101/013433 |doi= 10.1038/NATURE14317}}
• {{cite journal |ref={{sfnref|Haber et al.|2012}} |vauthors=Haber M, Platt DE, Ashrafian Bonab M, Youhanna SC, Soria-Hernanz DF, Martínez-Cruz B, Douaihy B, Ghassibe-Sabbagh M, Rafatpanah H, Ghanbari M, Whale J, Balanovsky O, Wells RS, Comas D, Tyler-Smith C, Zalloua PA |display-authors=8 |title=Afghanistan's ethnic groups share a Y-chromosomal heritage structured by historical events |journal=PLOS ONE |volume=7 |issue=3 |at=e34288 |year=2012 |pmid= 22470552 |pmc= 3314501 |doi= 10.1371/journal.pone.0034288 |bibcode= 2012PLoSO...734288H |doi-access=free }}
• {{cite web|author=ISOGG|year=2012|url=http://www.isogg.org/tree/ISOGG_HapgrpO.html|title=Y-DNA Haplogroup O and its Subclades - 2012}}
• {{Cite journal |ref={{sfnref|Karafet et al.|2014}} |last1=Karafet |first1=Tatiana M. |last2=Mendez |first2=Fernando L. |last3=Sudoyo |first3=Herawati |last4=Lansing |first4=J. Stephen |last5=Hammer |first5=Michael F. |title= Improved phylogenetic resolution and rapid diversification of Y-chromosome haplogroup K-M526 in Southeast Asia |journal=Nature |year=2014 |volume=23 |issue=3 |pages=369–373 |doi= 10.1038/ejhg.2014.106 |doi-access=free |pmid=24896152 |pmc=4326703}}
• {{Cite journal |last1=Kasperaviciūte |first1=D. |last2=Kucinskas |first2=V. |last3=Stoneking |first3=M. |s2cid=26562505 |year=2005 |title=Y Chromosome and Mitochondrial DNA Variation in Lithuanians |journal=Annals of Human Genetics |volume=68 |issue=5 |pages=438–452 |doi=10.1046/j.1529-8817.2003.00119.x |pmid=15469421 }}
• {{Cite journal |ref={{sfnref|Kayser et al.|2005}} |vauthors=Kayser M, Lao O, Anslinger K, Augustin C, Bargel G, Edelmann J, Elias S, Heinrich M, Henke J, Henke L, Hohoff C, Illing A, Jonkisz A, Kuzniar P, Lebioda A, Lessig R, Lewicki S, MacIejewska A, Monies DM, Pawłowski R, Poetsch M, Schmid D, Schmidt U, Schneider PM, Stradmann-Bellinghausen B, Szibor R, Wegener R, Wozniak M, Zoledziewska M, Roewer L |s2cid=11066186 |title=Significant genetic differentiation between Poland and Germany follows present-day political borders, as revealed by Y-chromosome analysis |journal=Human Genetics |volume=117 |issue=5 |pages=428–443 |year=2005 |pmid=15959808 |doi=10.1007/s00439-005-1333-9 |display-authors=8 |url=http://dirkschweitzer.net/E3b-papers/HG-05-428-Poland-Germany.pdf |archive-url=https://web.archive.org/web/20090304100323/http://dirkschweitzer.net/E3b-papers/HG-05-428-Poland-Germany.pdf |archive-date=March 4, 2009 }}
• {{Cite journal |ref={{sfnref|Keyser et al.|2009}} |last1=Keyser |first1=Christine |last2=Bouakaze |first2=Caroline |last3=Crubézy |first3=Eric |last4=Nikolaev |first4=Valery G. |last5=Montagnon |first5=Daniel |last6=Reis |first6=Tatiana |last7=Ludes |first7=Bertrand |s2cid=21347353 |year=2009 |title=Ancient DNA provides new insights into the history of south Siberian Kurgan people |doi=10.1007/s00439-009-0683-0 |journal=Human Genetics |volume=126 |issue=3 |pages=395–410 |pmid=19449030}}
• {{cite journal |ref={{sfnref|Kim et al.|2010}} |last1=Kim |first1=Kijeong |last2=Brenner |first2=Charles H. |last3=Mair |first3=Victor H. |last4=Lee |first4=Kwang-Ho |last5=Kim |first5=Jae-Hyun |last6=Gelegdorj |first6=Eregzen |last7=Batbold |first7=Natsag |last8=Song |first8=Yi-Chung |last9=Yun |first9=Hyeung-Won |last10=Chang |first10=Eun-Jeong |last11=Lkhagvasuren |first11=Gavaachimed |last12=Bazarragchaa |first12=Munkhtsetseg |last13=Park |first13=Ae-Ja |last14=Lim |first14=Inja |last15=Hong |first15=Yun-Pyo |last16=Kim |first16=Wonyong |last17=Chung |first17=Sang-In |last18=Kim |first18=Dae-Jin |last19=Chung |first19=Yoon-Hee |last20=Kim |first20=Sung-Su |last21=Lee |first21=Won-Bok |last22=Kim |first22=Kyung-Yong |display-authors=8 |title=A western Eurasian male is found in 2000-year-old elite Xiongnu cemetery in Northeast Mongolia |journal=American Journal of Physical Anthropology |volume=142 |issue=3 |year=2010 |pages=429–440 |doi= 10.1002/ajpa.21242 |pmid= 20091844}}
• {{Cite journal |ref={{sfnref|Kivisild et al.|2003}} |last1=Kivisild |first1=T |year=2003 |title=The Genetic Heritage of the Earliest Settlers Persists Both in Indian Tribal and Caste Populations |periodical=AJHG |volume=72 |issue=2 |doi=10.1086/346068 |pmc=379225 |pmid=12536373 |pages=313–32 |display-authors=8 |last2=Rootsi |first2=S |last3=Metspalu |first3=M |last4=Mastana |first4=S |last5=Kaldma |first5=K |last6=Parik |first6=J |last7=Metspalu |first7=E |last8=Adojaan |first8=M |last9=Tolk |first9=HV |last10=Stepanov |first10=V. |last11=Gölge |first11=M. |last12=Usanga |first12=E. |last13=Papiha |first13=S.S. |last14=Cinnioğlu |first14=C. |last15=King |first15=R. |last16=Cavalli-Sforza |first16=L. |last17=Underhill |first17=P.A. |last18=Villems |first18=R. }}
• {{cite journal |last=Lazaridis |first=Iosif |display-authors=etal |year=2016 |title=Genomic insights into the origin of farming in the ancient Near East |journal=Nature |volume=536 |issue=7617 |pages=419–424 |doi= 10.1038/nature19310 |pmid=27459054 |pmc=5003663 |bibcode=2016Natur.536..419L |ref={{sfnref |Lazaridis et al. |2016}} }}
• {{Cite journal |vauthors=Lell JT, Sukernik RI, Starikovskaya YB, Su B, Jin L, Schurr TG, Underhill PA, Wallace DC |title=The Dual Origin and Siberian Affinities of Native American Y Chromosomes |journal=American Journal of Human Genetics |volume=70 |issue=1 |pages=192–206 |year=2002 |ref={{sfnref|Lell et al.|2002}} |url=http://hpgl.stanford.edu/publications/AJHG_2002_v70_p192-206.pdf |url-status=dead |archive-url=https://web.archive.org/web/20030422095817/http://hpgl.stanford.edu/publications/AJHG_2002_v70_p192-206.pdf |archive-date=April 22, 2003 |doi=10.1086/338457 |pmc=384887 |pmid=11731934 }}
• {{cite journal |ref={{sfnref|Liu Shu-hu et al.|2018}} |author=Liu Shu-hu |author2=Nizam Yilihamu |author3=Rabiyamu Bake |author4=Abdukeram Bupatima |author5=Dolkun Matyusup |date=2018 |title=A study of genetic diversity of three isolated populations in Xinjiang using Y-SNP |journal=Acta Anthropologica Sinica |volume=37 |issue=1 |pages=146–156}}
• {{cite news |author=Carlos Quiles |date=September 10, 2018 |title=A study of genetic diversity of three isolated populations in Xinjiang using Y-SNP |newspaper=Indo-European.eu |url=https://indo-european.eu/2018/09/a-study-of-genetic-diversity-of-three-isolated-populations-in-xinjiang-using-y-snp/ |access-date=July 24, 2020 |archive-date=September 20, 2020 |archive-url=https://web.archive.org/web/20200920145543/https://indo-european.eu/2018/09/a-study-of-genetic-diversity-of-three-isolated-populations-in-xinjiang-using-y-snp/ |url-status=live }}

• {{cite journal |ref={{sfnref|Malmström et al.|2019}} |last1=Malmström |first1=Helena |last2=Günther |first2=Torsten |last3=Svensson |first3=Emma M. |last4=Juras |first4=Anna |last5=Fraser |first5=Magdalena |last6=Munters |first6=Arielle R. |last7=Pospieszny |first7=Łukasz |last8=Tõrv |first8=Mari |last9=Lindström |first9=Jonathan |display-authors=8 |date=October 9, 2019 |title=The genomic ancestry of the Scandinavian Battle Axe Culture people and their relation to the broader Corded Ware horizon |journal=Proceedings of the Royal Society B |volume=286 |issue=1912 |doi=10.1098/rspb.2019.1528 |doi-access=free |pmid= 31594508|pmc=6790770 }}
• {{cite journal |ref={{sfnref|Malyarchuk et al.|2013}} |last1=Malyarchuk |first1=Boris |last2=Derenko |first2=Miroslava |last3=Wozniak |first3=Marcin |last4=Grzybowski |first4=Tomasz |s2cid=2752490 |year=2013 |title=Y-chromosome variation in Tajiks and Iranians |journal=Annals of Human Biology |volume=40| issue=1| pages=48–54| doi= 10.3109/03014460.2012.747628 |pmid= 23198991 }}
• {{Cite journal |ref={{sfnref|Mascarenhas et al.|2015}} |last1=Mascarenhas |first1=Desmond D. |last2=Raina |first2=Anupuma |last3=Aston |first3=Christopher E. |last4=Sanghera |first4=Dharambir K. |year=2015 |title=Genetic and Cultural Reconstruction of the Migration of an Ancient Lineage |journal=BioMed Research International |volume=2015 |page=651415 |doi=10.1155/2015/651415 |pmid=26491681 |pmc=4605215 |doi-access=free }}
• {{Cite bioRxiv |ref={{sfnref|Mathieson et al.|2015}} |last1= Mathieson |first1= Iain |last2= Lazaridis |first2= Iosif |last3= Rohland |first3= Nadin |last4= Mallick |first4= Swapan |last5= Patterson |first5= Nick |last6= Alpaslan Roodenberg |first6= Songul |last7= Harney |first7= Eadaoin |last8= Stewardson |first8= Kristin |last9= Fernandes |first9= Daniel |last10= Novak |first10= Mario |last11= Sirak |first11= Kendra |last12= Gamba |first12= Cristina |last13= Jones |first13= Eppie R. |last14= Llamas |first14= Bastien |last15= Dryomov |first15= Stanislav |last16= Pickrell |first16= Joseph |last17= Arsuaga |first17= Juan Luis |last18= De Castro |first18= Jose Maria Bermudez |last19= Carbonell |first19= Eudald |last20= Gerritsen |first20= Fokke |last21= Khokhlov |first21= Aleksandr |last22= Kuznetsov |first22= Pavel |last23= Lozano |first23= Marina |last24= Meller |first24= Harald |last25= Mochalov |first25= Oleg |last26= Moiseyev |first26= Vayacheslav |last27= Rojo Guerra |first27= Manuel A. |last28= Roodenberg |first28= Jacob |last29= Verges |first29= Josep Maria |last30= Krause |first30= Johannes |display-authors=8 |title= Eight thousand years of natural selection in Europe |year=2015 |biorxiv= 10.1101/016477}}
• {{Cite journal |last1=Mirabal |first1=Sheyla |year=2009 |title=Y-Chromosome distribution within the geo-linguistic landscape of northwestern Russia |journal=European Journal of Human Genetics |pmid=19259129 |volume=17 |issue=10 |pmc=2986641 |pages=1260–1273 |doi=10.1038/ejhg.2009.6 |last2=Regueiro |first2=M |last3=Cadenas |first3=AM |last4=Cavalli-Sforza |first4=LL |last5=Underhill |first5=PA |last6=Verbenko |first6=DA |last7=Limborska |first7=SA |last8=Herrera |first8=RJ |display-authors=etal |ref={{harvid |Mirabal et al. |2009}} }}
• {{cite journal |ref={{sfnref|Mohammad et al.|2009}} |vauthors=Mohammad T, Xue Y, Evison M, Tyler-Smith C |date=2009 |title=Genetic structure of nomadic Bedouin from Kuwait |journal=Heredity |volume=103 |issue=5 |pages=425–433 |doi= 10.1038/hdy.2009.72 |pmid=19639002 |pmc=2869035 |doi-access=free}}

• {{cite bioRxiv | last1 =Narasimhan | first1 =Vagheesh M. | last2 =Anthony | first2 =David | last3 =Mallory | first3 =James | last4 =Reich | first4 =David | year =2018 | title =The Genomic Formation of South and Central Asia | biorxiv =10.1101/292581 |ref={{sfnref|Narasimhan et al.|2018}}}}
• {{Citation | ref={{sfnref|Narasimhan et al.|2019}} | last1 =Narasimhan | first1 =Vagheesh M. | year =2019 | last2 =Patterson | first2 = N.J. |last3=Moorjani|first3=Priya|last4=Rohland|first4=Nadin| display-authors =etal | title =The Formation of Human Populations in South and Central Asia | journal =Science | volume =365 | issue =6457 | pages =eaat7487 | url =https://scholar.harvard.edu/vagheesh/centralsouthasia| doi =10.1126/science.aat7487 | pmid =31488661 | pmc =6822619 }}
• {{Cite journal |ref={{sfnref|Nasidze et al.|2004}} |vauthors=Nasidze I, Ling EY, Quinque D, Dupanloup I, Cordaux R, Rychkov S, Naumova O, Zhukova O, Sarraf-Zadegan N, Naderi GA, Asgary S, Sardas S, Farhud DD, Sarkisian T, Asadov C, Kerimov A, Stoneking M |display-authors=8 |title=Mitochondrial DNA and Y-Chromosome Variation in the Caucasus |journal=Annals of Human Genetics |volume=68 |issue=Pt 3 |pages=205–221 |year=2004 |pmid=15180701 |doi=10.1046/j.1529-8817.2004.00092.x |s2cid=27204150 |url=http://www.eva.mpg.de/genetics/pdf/Caucasus_big_paper.pdf |url-status=dead |archive-url=https://web.archive.org/web/20041030074136/http://www.eva.mpg.de/genetics/pdf/Caucasus_big_paper.pdf |archive-date=October 30, 2004 }}
• {{Cite journal |ref={{sfnref|Nasidze et al.|2005}} |vauthors=Nasidze I, Quinque D, Ozturk M, Bendukidze N, Stoneking M |title=MtDNA and Y-chromosome Variation in Kurdish Groups |year=2005 |journal=Annals of Human Genetics |volume=69 |issue=Pt 4 |pages=401–412 |pmid=15996169 |doi=10.1046/j.1529-8817.2005.00174.x |s2cid=23771698 |url=http://www.eva.mpg.de/genetics/pdf/Kurds.pdf |archive-url=https://web.archive.org/web/20090823162702/http://www.eva.mpg.de/genetics/pdf/Kurds.pdf |archive-date=August 23, 2009 |url-status=dead }}
• {{Citation |ref={{sfnref|Pamjav et al.|2012}} |last1=Pamjav |first1=Horolma |first2=Tibor |last2=Fehér |first3=Endre |last3=Németh |first4=Zsolt |last4=Pádár |s2cid=4820868 |year=2012 |title=Brief communication: new Y-chromosome binary markers improve phylogenetic resolution within haplogroup R1a1 |journal=American Journal of Physical Anthropology |volume=149 |issue=4 |pages=611–615 |doi=10.1002/ajpa.22167 |pmid=23115110}}
• {{Cite journal |vauthors=Passarino G, Cavalleri GL, Lin AA, Cavalli-Sforza LL, Børresen-Dale AL, Underhill |first6=PA |year=2002 |title=Different genetic components in the Norwegian population revealed by the analysis of mtDNA and Y chromosome polymorphisms |journal=European Journal of Human Genetics |volume=10 |issue=9 |pages=521–529 |pmid=12173029 |doi=10.1038/sj.ejhg.5200834 |doi-access=free |ref={{sfnref|Passarino et al.|2002}} }}
• {{cite journal |ref={{sfnref|Pathak et al.|2018}} |last1=Pathak |first1=Ajai K. |last2=Kadian |first2=Anurag |last3=Kushniarevich |first3=Alena |last4=Montinaro |first4=Francesco |last5=Mondal |first5=Mayukh |last6=Ongaro |first6=Linda |last7=Singh |first7=Manvendra |last8=Kumar |first8=Pramod |last9=Rai |first9=Niraj |last10=Parik |first10=Jüri |last11=Metspalu |first11=Ene |last12=Rootsi |first12=Siiri |last13=Pagani |first13=Luca |last14=Kivisild |first14=Toomas |last15=Metspalu |first15=Mait |last16=Chaubey |first16=Gyaneshwer |last17=Villems |first17=Richard |display-authors=8 |title=The Genetic Ancestry of Modern Indus Valley Populations from Northwest India |journal=The American Journal of Human Genetics |date=December 6, 2018 |volume=103 |issue=6 |pages=918–929 |doi=10.1016/j.ajhg.2018.10.022 |pmid=30526867 |pmc=6288199 }}
• {{Cite journal |ref={{sfnref|Pawlowski et al.|2002}} |last1=Pawlowski |first1=R |last2=Dettlaff-Kakol |first2=A |last3=MacIejewska |first3=A |last4=Paszkowska |first4=R |last5=Reichert |first5=M |last6=Jezierski |first6=G |title=Population genetics of 9 Y-chromosome STR loci w Northern Poland |journal=Arch. Med. Sadowej Kryminol. |volume=52 |issue=4 |pages=261–277 |year=2002 |pmid=14669672}}
• {{Cite journal |vauthors=Pericić M, Lauc LB, Klarić IM, Rootsi S, Janićijević B, Rudan I, Terzić R, Colak I, Kvesić A, Popović D, Sijacki A, Behluli I, Dordevic D, Efremovska L, Bajec DD, Stefanović BD, Villems R, Rudan P |year=2005 |title=High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations |journal=Mol. Biol. Evol. |volume=22 |issue=10 |pages=1964–75 |pmid=15944443 |doi=10.1093/molbev/msi185 |doi-access=free |display-authors=8 |ref={{sfnref|Pericić et al.|2005}} }}
• {{cite journal |ref={{sfnref|Poznik et al.|2016}} |vauthors=Poznik GD |display-authors=etal |year=2016 |title=Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences |journal=Nature Genetics |doi=10.1038/ng.3559 |volume=48 |issue=6 |pages=593–599 |pmid=27111036 |pmc=4884158 |hdl=11858/00-001M-0000-002A-F024-C}}
• {{Cite journal |last1=Regueiro |first1=M |year=2006 |title=Iran: Tricontinental Nexus for Y-Chromosome Driven Migration |journal=Hum Hered |volume=61 |issue=3 |pages=132–143 |doi=10.1159/000093774 |pmid=16770078 |last2=Cadenas |first2=AM |last3=Gayden |first3=T |last4=Underhill |first4=PA |last5=Herrera |first5=RJ |s2cid=7017701 |ref={{harvid |Regueiro |2006}}}}
• {{cite journal |ref={{sfnref|Ricaut et al.|2004}} |vauthors=Ricaut F, Keyser-Tracqui C, Bourgeois I, Crubézy E, Ludes B |s2cid=35948291 |year=2004 |title=Genetic Analysis of a Scytho-Siberian Skeleton and Its Implications for Ancient Central Asian Migrations |journal=Human Biology |volume=76 |issue= 1|pages=109–25 |doi= 10.1353/hub.2004.0025 |pmid= 15222683 }}
• {{Cite journal |vauthors=Rosser ZH, Zerjal T, Hurles ME, Adojaan M, Alavantic D, Amorim A, Amos W, Armenteros M, Arroyo E, Barbujani G, Beckman G, Beckman L, Bertranpetit J, Bosch E, Bradley DG, Brede G, Cooper G, Côrte-Real HB, De Knijff P, Decorte R, Dubrova YE, Evgrafov O, Gilissen A, Glisic S, Gölge M, Hill EW, Jeziorowska A, Kalaydjieva L, Kayser M, Kivisild T |title=Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language |journal=American Journal of Human Genetics |volume=67 |issue=6 |pages=1526–1543 |year=2000 |doi=10.1086/316890 |doi-access=free |pmc=1287948 |pmid=11078479 |display-authors=8 |ref={{sfnref|Rosser et al.|2000}} }}
• {{cite journal |last1=Saag |first1=Lehti |last2=Varul |first2=Liivi |last3=Scheib |first3=Christiana Lyn |last4=Stenderup |first4=Jesper |last5=Allentoft |first5=Morten E. |last6=Saag|first6=Lauri |last7=Pagani|first7=Luca |last8=Reidla |first8=Maere |last9=Tambets |first9=Kristiina |display-authors=8 |date=July 24, 2017 |title=Extensive Farming in Estonia Started through a Sex-Biased Migration from the Steppe |journal=Current Biology |publisher=Cell Press |volume=27 |issue=14 |pages=2185–2193 |doi=10.1016/j.cub.2017.06.022 |doi-access=free |pmid= 28712569 |bibcode=2017CBio...27E2185S |ref={{harvid|Saag et al.|2017}}}}
• {{cite bioRxiv |last1=Saag |first1=Lehti |last2=Vasilyev |first2=Sergey V. |last3=Varul |first3=Liivi |last4=Kosorukova |first4=Natalia V. |last5=Gerasimov |first5=Dmitri V. |last6=Oshibkina |first6=Svetlana V. |last7=Griffith |first7=Samuel J. |last8=Solnik |first8=Anu |last9=Saag|first9=Lauri |display-authors=8 |date=July 3, 2020 |title=Genetic ancestry changes in Stone to Bronze Age transition in the East European plain |biorxiv=10.1101/2020.07.02.184507 |ref={{harvid|Saag et al.|2020}}}}
• {{Cite journal |last1=Sahoo |first1=S |title=A prehistory of Indian Y chromosomes: Evaluating demic diffusion scenarios |journal=Proceedings of the National Academy of Sciences |volume=103 |issue=4 |pages=843–848 |year=2006 |doi=10.1073/pnas.0507714103 |pmc=1347984 |pmid=16415161 |bibcode=2006PNAS..103..843S |last2=Singh |first2=A |last3=Himabindu |first3=G |last4=Banerjee |first4=J |last5=Sitalaximi |first5=T |last6=Gaikwad |first6=S |last7=Trivedi |first7=R |last8=Endicott |first8=P |last9=Kivisild |first9=T |last10=Metspalu |first10=M. |last11=Villems |first11=R. |last12=Kashyap |first12=V. K. |display-authors=8 |ref={{harvid |Sahoo et al. |2006}}|doi-access=free }}
• {{Cite journal |vauthors=Scozzari R, Cruciani F, Pangrazio A, Santolamazza P, Vona G, Moral P, Latini V, Varesi L, Memmi MM, Romano V, De Leo G, Gennarelli M, Jaruzelska J, Villems R, Parik J, MacAulay V, Torroni A |year=2001 |title=Human Y-Chromosome Variation in the Western Mediterranean Area: Implications for the Peopling of the Region |url=http://evolutsioon.ut.ee/publications/Scozzari2001.pdf |journal=Human Immunology |volume=62 |issue=9 |doi=10.1016/S0198-8859(01)00286-5 |pages=871–84 |display-authors=8 |pmid=11543889 |citeseerx=10.1.1.408.4857 |ref={{sfnref|Scozzari et al.|2001}} |access-date=March 15, 2009 |archive-date=December 17, 2012 |archive-url=https://web.archive.org/web/20121217135021/http://evolutsioon.ut.ee/publications/Scozzari2001.pdf |url-status=dead }}
• {{Citation |last1=Semenov |first1=Alexander S. |last2=Bulat |first2=Vladimir V. |s2cid=172131289 |year=2016 |title=Ancient Paleo-DNA of Pre-Copper Age North-Eastern Europe: Establishing the Migration Traces of R1a1 Y-DNA Haplogroup |journal= European Journal of Molecular Biotechnology |volume=11 |issue=1 |pages=40–54 |doi= 10.13187/ejmb.2016.11.40 }}
• {{Cite journal |last1=Semino |first1=O |last2=Passarino |first2=G |last3=Oefner |first3=PJ |last4=Lin |first4=AA |last5=Arbuzova |first5=S |last6=Beckman |first6=LE |last7=De Benedictis |first7=G |last8=Francalacci |first8=P |last9=Kouvatsi |first9=A |last10=Limborska |first10=S |last11=Marcikiae |first11=M |last12=Mika |first12=A |last13=Mika |first13=B |last14=Primorac |first14=D |last15=Santachiara-Benerecetti |first15=A. S. |last16=Cavalli-Sforza |first16=L. L. |last17=Underhill |first17=P. A. |year=2000 |title=The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective |journal=Science |volume=290 |issue=5494 |pages=1155–1159 |pmid=11073453 |url=http://hpgl.stanford.edu/publications/Science_2000_v290_p1155.pdf |doi=10.1126/science.290.5494.1155 |bibcode=2000Sci...290.1155S |display-authors=8 |ref={{harvid |Semino et al.|2000}} |url-status=dead |archive-url=https://web.archive.org/web/20031125151213/http://hpgl.stanford.edu/publications/Science_2000_v290_p1155.pdf |archive-date=November 25, 2003 }}
• {{Cite journal |last1=Sengupta |year=2006 |title=Polarity and Temporality of High-Resolution Y-Chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influence of Central Asian Pastoralists |periodical=American Journal of Human Genetics |volume=78 |issue=2 |pages=202–21 |pmid=16400607 |pmc=1380230 |doi=10.1086/499411 |first1=S |last2=Zhivotovsky |first2=LA |last3=King |first3=R |last4=Mehdi |first4=SQ |last5=Edmonds |first5=CA |last6=Chow |first6=CE |last7=Lin |first7=AA |last8=Mitra |first8=M |last9=Sil |first9=SK |last10=Ramesh |first10=A. |last11=Usha Rani |first11=M.V. |last12=Thakur |first12=Chitra M. |last13=Cavalli-Sforza |first13=L. Luca |last14=Majumder |first14=Partha P. |last15=Underhill |first15=Peter A. |display-authors=8 |ref={{harvid |Sengupta |2006}}}}
• {{Cite journal |last1=Sharma |first1=S |last2=Rai |first2=E |last3=Sharma |first3=P |last4=Jena |first4=M |last5=Singh |first5=S |last6=Darvishi |first6=K |last7=Bhat |first7=AK |last8=Bhanwer |first8=AJ |last9=Tiwari |first9=PK |last10=Bamezai |first10=Rameshwar N K |title=The Indian origin of paternal haplogroup R1a1(*)substantiates the autochthonous origin of Brahmins and the caste system |journal=Journal of Human Genetics |volume=54 |issue=1 |pages=47–55 |year=2009 |doi=10.1038/jhg.2008.2 |pmid=19158816 |display-authors=8 |ref={{harvid |Sharma et al. |2009}}|doi-access=free }}
• {{Cite thesis |last=Schilz |first=Felix |title=Molekulargenetische Verwandtschaftsanalysen am prähistorischen Skelettkollektiv der Lichtensteinhöhle |type=Dissertation |publisher=Mathematisch-Naturwissenschaftlichen Fakultäten der Georg-August-Universität |location=Göttingen |year=2006 |url=http://webdoc.sub.gwdg.de/diss/2006/schilz/schilz.pdf |lang=de |trans-title=Molecular genetic kinship analysis on the prehistoric skeleton collective of the Lichtenstein Cave |access-date=November 9, 2009 |archive-date=June 8, 2011 |archive-url=https://web.archive.org/web/20110608140210/http://webdoc.sub.gwdg.de/diss/2006/schilz/schilz.pdf |url-status=live }}
• {{Cite journal |last1=Silva |first1=Marina |display-authors=etal |title=A genetic chronology for the Indian Subcontinent points to heavily sex-biased dispersals |journal=BMC Evolutionary Biology |year=2017 |page=88 |doi= 10.1186/s12862-017-0936-9 |pmid= 28335724 |pmc= 5364613 |volume=17 |issue=1 |bibcode=2017BMCEE..17...88S |ref={{harvid |Silva et al. |2017}} |doi-access=free }}
• {{Cite journal |vauthors=Tambets K, Rootsi S, Kivisild T, Help H, Serk P, Loogväli EL, Tolk HV, Reidla M, Metspalu E, Pliss L, Balanovsky O, Pshenichnov A, Balanovska E, Gubina M, Zhadanov S, Osipova L, Damba L, Voevoda M, Kutuev I, Bermisheva M, Khusnutdinova E, Gusar V, Grechanina E, Parik J, Pennarun E, Richard C, Chaventre A, Moisan JP, Barać L, Peričić M |title=The Western and Eastern Roots of the Saami—the Story of Genetic 'Outliers' Told by Mitochondrial DNA and Y Chromosomes |journal=American Journal of Human Genetics |volume=74 |issue=4 |pages=661–682 |year=2004 |url= |pmid=15024688 |pmc=1181943 |doi=10.1086/383203 |doi-access=free |display-authors=8 |ref={{sfnref|Tambets et al.|2004}} }}
• {{Cite journal |last1=Thangaraj |year=2010 |title=The Influence of Natural Barriers in Shaping the Genetic Structure of Maharashtra Populations |last2=Naidu |last3=Crivellaro |last4=Tamang |last5=Upadhyay |journal=PLOS ONE |volume=5 |issue=12 |pages=e15283 |pmid=21187967 |pmc=3004917 |doi=10.1371/journal.pone.0015283 |bibcode=2010PLoSO...515283T |editor1-last=Cordaux |editor1-first=Richard |first1=Kumarasamy |first2=B. Prathap |first3=Federica |first4=Rakesh |first5=Shashank |last6=Sharma |first6=Varun Kumar |last7=Reddy |first7=Alla G. |last8=Walimbe |first8=S. R. |last9=Chaubey |first9=Gyaneshwer |last10=Kivisild |first10=T |last11=Singh |first11=L |display-authors=8 |ref={{harvid |Thangaraj et al. |2010}}|doi-access=free }}
• {{Cite journal |ref={{harvid |Underhill et al.|2009}} |last1=Underhill |first1=PA |last2=Myres |first2=NM |last3=Rootsi |first3=S |last4=Metspalu |first4=M |last5=Zhivotovsky |first5=LA |last6=King |first6=RJ |last7=Lin |first7=AA |last8=Chow |first8=CE |last9=Semino |first9=O |last10=Battaglia |first10=V |last11=Kutuev |first11=I |last12=Järve |first12=M |last13=Chaubey |first13=G |last14=Ayub |first14=Q |last15=Mohyuddin |first15=A |last16=Mehdi |first16=SQ |last17=Sengupta |first17=S |last18=Rogaev |first18=EI |last19=Khusnutdinova |first19=EK |last20=Pshenichnov |first20=A |last21=Balanovsky |first21=O |last22=Balanovska |first22=E |last23=Jeran |first23=N |last24=Augustin |first24=DH |last25=Baldovic |first25=M |last26=Herrera |first26=RJ |last27=Thangaraj |first27=K |last28=Singh |first28=V |last29=Singh |first29=L |last30=Majumder |first30=P |last31=Rudan |first31=P |last32=Primorac |first32=D |last33=Villems |first33=R |last34=Kivisild |first34=T |date=November 4, 2009 |publication-date=April 2010 |title=Separating the post-Glacial coancestry of European and Asian Y chromosomes within haplogroup R1a |volume=18 |issue=4 |journal=European Journal of Human Genetics |doi=10.1038/ejhg.2009.194 |doi-access=free |pmid=19888303 |pmc=2987245 |pages=479–84 |display-authors=8}}
• {{cite journal |ref={{sfnref|Underhill et al.|2014}} |last1=Underhill |first1=Peter A. |display-authors=etal |date=March 26, 2014 |publication-date=January 2015 |title=The phylogenetic and geographic structure of Y-chromosome haplogroup R1a |journal=European Journal of Human Genetics |volume=23 |issue=1 |pages=124–131 |doi=10.1038/ejhg.2014.50 |doi-access=free |pmid=24667786 |pmc=4266736}} {{Cite web |access-date=June 12, 2016 |url=http://thebigone.stanford.edu/papers/Underhill_phylogenetic_March-2014.pdf |archive-url=https://web.archive.org/web/20160816180616/http://thebigone.stanford.edu/papers/Underhill_phylogenetic_March-2014.pdf |archive-date=August 16, 2016 |url-status=dead |title=PDF }}
• {{Cite journal |ref={{sfnref|Wang et al.|2003}} |last1=Wang |first1=Wei |last2=Wise |first2=Cheryl |last3=Baric |first3=Tom |last4=Black |first4=Michael L. |last5=Bittles |first5=Alan H. |s2cid=11138499 |year=2003 |doi=10.1007/s00439-003-0948-y |title=The origins and genetic structure of three co-resident Chinese Muslim populations: The Salar, Bo'an and Dongxiang |journal=Human Genetics |volume=113 |issue=3 |pages=244–52 |pmid=12759817}}
• {{Citation |last=Wells |first=R.S. |year=2001 |title=The Eurasian Heartland: A continental perspective on Y-chromosome diversity |journal=Proceedings of the National Academy of Sciences of the USA |volume=98 |issue=18 |pages=10244–10249 |doi=10.1073/pnas.171305098 |pmid=11526236 |pmc=56946|bibcode=2001PNAS...9810244W |doi-access=free }}
• {{Cite journal |last1=Yan |first1=Shi |last2=Wang |first2=Chuan-Chao |last3=Zheng |first3=Hong-Xiang |last4=Wang |first4=Wei |last5=Qin |first5=Zhen-Dong |last6=Wei |first6=Lan-Hai |last7=Wang |first7=Yi |last8=Pan |first8=Xue-Dong |last9=Fu |first9=Wen-Qing |last10=He |first10=Yun-Gang |last11=Xiong |first11=Li-Jun |last12=Jin |first12=Wen-Fei |last13=Li |first13=Shi-Lin |last14=An |first14=Yu |last15=Li |first15=Hui |last16=Jin |first16=Li |last17=Su |first17=Bing |display-authors=8 |title=Y Chromosomes of 40% Chinese Descend from Three Neolithic Super-Grandfathers |journal=PLOS ONE |date=August 29, 2014 |volume=9 |issue=8 |at=e105691 |doi= 10.1371/journal.pone.0105691 |doi-access=free |pmid= 25170956 |pmc= 4149484 |arxiv=1310.3897 |bibcode= 2014PLoSO...9j5691Y |ref={{sfnref|Yan et al.|2014}} }}
• {{Cite journal |ref=CITEREFYCC2002 |doi=10.1101/gr.217602 |author=Y Chromosome Consortium "YCC" |year=2002 |title=A Nomenclature System for the Tree of Human Y-Chromosomal Binary Haplogroups |periodical=Genome Research |volume=12 |pages=339–348 |issue=2 |pmid=11827954 |pmc=155271}}
• {{cite journal |ref={{sfnref|Zerjal et al.|2002}} |doi=10.1086/342096 |title=A Genetic Landscape Reshaped by Recent Events: Y-Chromosomal Insights into Central Asia |year=2002 |last1=Zerjal |first1=Tatiana |last2=Wells |first2=R. Spencer |last3=Yuldasheva |first3=Nadira |last4=Ruzibakiev |first4=Ruslan |last5=Tyler-Smith |first5=Chris |journal=The American Journal of Human Genetics |volume=71 |issue=3 |pages=466–82 |pmid=12145751 |pmc=419996}}
• {{cite journal |ref={{sfnref|Zhong et al.|2011}} |vauthors=Zhong H, Shi H, Qi XB, Duan Y, Tan PP, Jin L, SU B, Ma RZ |title=Extended Y chromosome investigation suggests postglacial migrations of modern humans into East Asia via the northern route |journal=Molecular Biology and Evolution |volume=28 |issue=1 |pages=717–27 |date=January 2011 |pmid=20837606 |doi=10.1093/molbev/msq247 |doi-access=free }}
• {{Cite journal |ref={{sfnref|Zhou et al.|2007}} |last1=Zhou |first1=Ruixia |last2=An |first2=Lizhe |last3=Wang |first3=Xunling |last4=Shao |first4=Wei |last5=Lin |first5=Gonghua |last6=Yu |first6=Weiping |last7=Yi |first7=Lin |last8=Xu |first8=Shijian |last9=Xu |first9=Jiujin |last10=Xie |first10=Xiaodong |display-authors=8 |title=Testing the hypothesis of an ancient Roman soldier origin of the Liqian people in northwest China: a Y-chromosome perspective |journal=Journal of Human Genetics |volume=52 |issue=7 |year=2007 |doi=10.1007/s10038-007-0155-0 |pmid=17579807 |pages=584–91 |doi-access=free }}

{{refend}}

{{refbegin}}

• {{Cite book |author=Gimbutas |title=Indo-European and Indo-Europeans |publisher=Univ. of Pennsylvania Press |location=Philadelphia, PA |year=1970 |pages=155–195}}
• {{Cite book |author=Patrizia Malaspina |author2=Andrey I. Kozlov |author3=Fulvio Cruciani |author4=Piero Santolamazza |author5=Nejat Akar |author6=Dimiter Kovatchev |author7=Marina G. Kerimova |author8=Juri Parik |author9=Richard Villems |author10=Rosana Scozzari |author11=Andrea Novelletto |year=2003 |chapter=Analysis of Y-chromosome variation in modern populations at the European-Asian border |pages=309–313 |chapter-url=http://evolutsioon.ut.ee/publications/Malaspina2003.pdf |editor=K. Boyle |editor2=C. Renfrew |editor3=M. Levine |title=Ancient interactions: east and west in Eurasia |series=McDonald Institute Monographs |publisher=Cambridge University Press |location=Cambridge |access-date=September 10, 2009 |archive-date=June 24, 2023 |archive-url=https://web.archive.org/web/20230624125920/http://evolutsioon.ut.ee/publications/Malaspina2003.pdf |url-status=dead }}
• {{cite book |last=Parpola |first=Asko |year=2015 |title=The Roots of Hinduism. The early Aryans and the Indus Civilisation |publisher=Oxford University Press}}
• {{Cite book |last=Sharma |first=S. |section=1344/T: The Autochthonous Origin and a Tribal Link of Indian Brahmins: Evaluation Through Molecular Genetic Markers |type=Abstract |title=The American Society of Human Genetics 57th Annual Meeting October 23–27, 2007; San Diego, California |page=273 |year=2007 |url=http://www.ashg.org/genetics/ashg/annmeet/2007/call/abstractbook.pdf |archive-url=https://web.archive.org/web/20080626010746/http://www.ashg.org/genetics/ashg/annmeet/2007/call/abstractbook.pdf |archive-date=June 26, 2008 }}
• {{cite book |last=Trautmann |first=Thomas |year=2005 |title=The Aryan Debate |publisher=Oxford University Press }}
• {{Cite thesis |last=Varzari |first=Alexander |year=2006 |title=Population History of the Dniester-Carpathians: Evidence from Alu Insertion and Y-Chromosome Polymorphisms |type=Dissertation |publisher=Ludwig-Maximilians-Universität |location=München |url=http://edoc.ub.uni-muenchen.de/5868/1/Varzari_Alexander.pdf |access-date=October 30, 2008 |archive-date=October 28, 2008 |archive-url=https://web.archive.org/web/20081028221059/http://edoc.ub.uni-muenchen.de/5868/1/Varzari_Alexander.pdf |url-status=live }}
• {{Cite book |last=Wells |first=Spencer |author-link=Spencer Wells |year=2002 |title=The Journey of Man: A Genetic Odyssey |publisher=Princeton University Press |isbn=978-0-691-11532-0 |url-access=registration |url=https://archive.org/details/journeyofmangene00well }}

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• {{Cite journal |last1=Adams |first1=Susan M. |last2=Bosch |first2=E |last3=Balaresque |first3=PL |last4=Ballereau |first4=SJ |last5=Lee |first5=AC |last6=Arroyo |first6=E |last7=López-Parra |first7=AM |last8=Aler |first8=M |last9=Grifo |first9=MS |last10=Brion |first10=Maria |last11=Carracedo |first11=Angel |last12=Lavinha |first12=João |last13=Martínez-Jarreta |first13=Begoña |last14=Quintana-Murci |first14=Lluis |last15=Picornell |first15=Antònia |last16=Ramon |first16=Misericordia |last17=Skorecki |first17=Karl |last18=Behar |first18=Doron M. |last19=Calafell |first19=Francesc |last20=Jobling |first20=Mark A. |title=The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula |journal=American Journal of Human Genetics |year=2008 |doi=10.1016/j.ajhg.2008.11.007 |volume=83 |pmc=2668061 |url=|issue=6 |pmid=19061982 |pages=725–36 |display-authors=8}}
• {{Cite journal |last1=Al Zahery |year=2003 |first1=N. |pages=458–72 |last2=Semino |first2=O. |last3=Benuzzi |first3=G. |last4=Magri |first4=C. |last5=Passarino |first5=G. |last6=Torroni |first6=A. |last7=Santachiara-Benerecetti |first7=A.S. |title=Y-chromosome and mtDNA polymorphisms in Iraq, a crossroad of the early human dispersal and of post-Neolithic migrations |issue=3 |url=http://www.familytreedna.com/pdf/Al_Zahery.pdf |volume=28 |doi=10.1016/S1055-7903(03)00039-3 |journal=Molecular Phylogenetics and Evolution |pmid=12927131 |bibcode=2003MolPE..28..458A |access-date=September 12, 2009 |archive-url=https://web.archive.org/web/20101227053418/http://www.familytreedna.com/pdf/Al_Zahery.pdf |archive-date=December 27, 2010 |url-status=dead }}
• {{Cite journal |last1=Bamshad |year=2001 |first1=M. |last2=Kivisild |first2=T |last3=Watkins |first3=WS |last4=Dixon |first4=ME |last5=Ricker |first5=CE |last6=Rao |first6=BB |last7=Naidu |first7=JM |last8=Prasad |first8=BV |last9=Reddy |first9=PG |last10=Rasanayagam |first10=A |last11=Papiha |first11=S. S. |last12=Villems |first12=R |last13=Redd |first13=A. J. |last14=Hammer |first14=M. F. |last15=Nguyen |first15=S. V. |last16=Carroll |first16=M. L. |last17=Batzer |first17=M. A. |last18=Jorde |first18=L. B. |title=Genetic evidence on the origins of Indian caste populations |journal=Genome Research |volume=11 |issue=6 |pages=994–1004 |pmid=11381027 |pmc=311057 |doi=10.1101/gr.GR-1733RR |display-authors=8}}
• {{Cite journal |last1=Barać |first1=Lovorka |last2=Pericić |first2=Marijana |last3=Klarić |first3=Irena Martinović |last4=Rootsi |first4=Siiri |last5=Janićijević |first5=Branka |last6=Kivisild |first6=Toomas |last7=Parik |first7=Jüri |last8=Rudan |first8=Igor |last9=Villems |first9=Richard |last10=Rudan |first10=Pavao |s2cid=15822710 |title=Y chromosomal heritage of Croatian population and its island isolates |journal=European Journal of Human Genetics |volume=11 |issue=7 |pages=535–42 |date=July 2003 |pmid=12825075 |doi=10.1038/sj.ejhg.5200992 |display-authors=8 |doi-access=free }}
• {{Cite journal |doi=10.1007/s00414-007-0177-3 |last1=Bouakaze |first1=C. |last2=Keyser |first2=C |last3=Amory |first3=S |last4=Crubézy |first4=E |last5=Ludes |first5=B |s2cid=22169008 |title=First successful assay of Y-SNP typing by SNaPshot minisequencing on ancient DNA |journal=International Journal of Legal Medicine |volume=121 |issue=6 |year=2007 |pages=493–9 |pmid=17534642}}
• {{Cite journal |title=Signatures of founder effects, admixture, and selection in the Ashkenazi Jewish population |first1=Steven |last1=Braya |first2=Jennifer |last2=Mullea |first3=Anne |last3=Dodda |first4=Ann |last4=Pulver |first5=Stephen |last5=Wooding |first6=Stephen |last6=Warren |journal=PNAS |volume=107 |issue=37 |pages=16222–16227 |year=2010 |pmid=20798349 |pmc=2941333 |doi=10.1073/pnas.1004381107 |bibcode=2010PNAS..10716222B|doi-access=free }}
• {{Cite journal |last1=Cinnioğlu |first1=C |last2=King |first2=R |last3=Kivisild |first3=T |last4=Kalfoğlu |first4=E |last5=Atasoy |first5=S |last6=Cavalleri |first6=GL |last7=Lillie |first7=AS |last8=Roseman |first8=CC |last9=Lin |first9=AA |last10=Prince |first10=Kristina |last11=Oefner |first11=Peter J. |last12=Shen |first12=Peidong |last13=Semino |first13=Ornella |last14=Cavalli-Sforza |first14=L. Luca |last15=Underhill |first15=Peter A. |s2cid=10763736 |title=Excavating Y-chromosome haplotype strata in Anatolia |journal=Hum Genet |volume=114 |year=2004 |issue=2 |doi=10.1007/s00439-003-1031-4 |pmid=14586639 |pages=127–48 |display-authors=8}}
• {{Cite journal |last1=Cordaux |year=2004 |first1=Richard |last2=Aunger |first2=R |last3=Bentley |first3=G |last4=Nasidze |first4=I |last5=Sirajuddin |first5=SM |last6=Stoneking |first6=M |s2cid=5721248 |title=Independent Origins of Indian Caste and Tribal Paternal Lineages |journal=Current Biology |pmid=14761656 |volume=14 |issue=3 |pages=231–235 |doi=10.1016/j.cub.2004.01.024|doi-access=free |bibcode=2004CBio...14..231C }}
• {{Cite journal |last1=Firasat |year=2006 |first1=Sadaf |last2=Khaliq |first2=S |last3=Mohyuddin |first3=A |last4=Papaioannou |first4=M |last5=Tyler-Smith |first5=C |last6=Underhill |first6=PA |last7=Ayub |first7=Q |title=Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan |pmid=17047675 |pmc=2588664 |doi=10.1038/sj.ejhg.5201726 |journal=European Journal of Human Genetics |volume=15 |issue=1 |pages=121–126}}
• {{Cite journal |last1=Flores |year=2005 |first1=Carlos |last2=Maca-Meyer |first2=N |last3=Larruga |first3=JM |last4=Cabrera |first4=VM |last5=Karadsheh |first5=N |last6=Gonzalez |first6=AM |title=Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan |journal=Journal of Human Genetics |volume=50 |issue=9 |pages=435–441 |doi=10.1007/s10038-005-0274-4 |pmid=16142507|doi-access=free }}
• {{Cite journal |last1=Hammer |year=2009 |first1=Michael F. |journal=Human Genetics |volume=126 |issue=5 |last2=Behar |pages=725–726 |doi=10.1007/s00439-009-0747-1 |first2=Doron M. |last3=Karafet |first3=Tatiana M. |last4=Mendez |first4=Fernando L. |last5=Hallmark |first5=Brian |last6=Erez |first6=Tamar |last7=Zhivotovsky |first7=Lev A. |last8=Rosset |first8=Saharon |last9=Skorecki |first9=Karl |title=Response|s2cid=264023342 }}
• {{Cite journal |last1=Helgason |first1=A |last2=Sigureardottir |year=2000 |first2=S |last3=Nicholson |first3=J |last4=Sykes |first4=B |last5=Hill |first5=E |last6=Bradley |issue=3 |first6=D |last7=Bosnes |first7=V |last8=Gulcher |first8=J |last9=Ward |first9=R |last10=Stefánsson |first10=Kári |title=Estimating Scandinavian and Gaelic Ancestry in the Male Settlers of Iceland |volume=67 |journal=American Journal of Human Genetics |doi=10.1086/303046 |pmid=10931763 |pmc=1287529 |pages=697–717 |display-authors=8}}
• {{cite journal |last1=Karafet |first1=T. M. |last2=Mendez |first2=F. L. |last3=Meilerman |first3=M. B. |last4=Underhill |first4=P. A. |last5=Zegura |first5=S. L. |last6=Hammer |first6=M. F. |url=http://genome.cshlp.org/content/early/2008/04/02/gr.7172008.abstract |title=New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree |journal=Genome Research |volume=18 |issue=5 |year=2008 |pages=830–838 |doi=10.1101/gr.7172008 |pmid=18385274 |pmc=2336805 |access-date=June 15, 2016 |archive-date=March 31, 2016 |archive-url=https://web.archive.org/web/20160331150544/http://genome.cshlp.org/content/early/2008/04/02/gr.7172008.abstract |url-status=live }} See also [http://www.genome.org/cgi/data/gr.7172008/DC1/1 Supplementary Material] {{Webarchive|url=https://web.archive.org/web/20080517231627/http://www.genome.org/cgi/data/gr.7172008/DC1/1 |date=May 17, 2008 }}.
• {{Cite journal |last1=Kharkov |first1=V. N. |last2=Stepanov |first2=V. A. |last3=Borinskaya |first3=S. A. |last4=Kozhekbaeva |first4=Zh. M. |last5=Gusar |first5=V. A. |last6=Grechanina |first6=E. Ya. |last7=Puzyrev |first7=V. P. |last8=Khusnutdinova |first8=E. K. |last9=Yankovsky |first9=N. K. |s2cid=25907265 |title=Gene Pool Structure of Eastern Ukrainians as Inferred from the Y-Chromosome Haplogroups |journal=Russian Journal of Genetics |volume=40 |issue=3 |pages=326–331 |year=2004 |doi=10.1023/B:RUGE.0000021635.80528.2f }} A copy can be found here [https://web.archive.org/web/20090304100322/http://dirkschweitzer.net/E3b-papers/Kharkov-RJG-2004-40-3-326.pdf ].
• {{Cite journal |last1=Kharkov |first1=V. N. |last2=Stepanov |first2=V. A. |last3=Feshchenko |first3=S. P. |last4=Borinskaya |first4=S. A. |last5=Yankovsky |first5=N. K. |last6=Puzyrev |first6=V. P. |s2cid=1357824 |title=Frequencies of Y Chromosome Binary Haplogroups in Belarusians |journal=Russian Journal of Genetics |volume=41 |issue=8 |pages=928–931 |year=2005 |doi=10.1007/s11177-005-0182-x |pmid=16161635 }} A copy can be found here [https://web.archive.org/web/20090304100321/http://dirkschweitzer.net/E3b-papers/Kharkov-RJG-2005-41-8-928.pdf ].
• {{Cite journal |last1=Kharkov |year=2007 |first1=V. N. |last2=Stepanov |journal=Russian Journal of Genetics |first2=V. A. |last3=Medvedeva |first3=O. F. |last4=Spiridonova |first4=M. G. |last5=Voevoda |first5=M. I. |last6=Tadinova |first6=V. N. |last7=Puzyrev |first7=V. P. |s2cid=566825 |title=Gene Pool Differences between Northern and Southern Altaians Inferred from the Data on Y-Chromosomal Haplogroups |pages=551–562 |url=http://www.medgenetics.ru/UserFile/File/Doc/Evolution%20Doc/Kharkov%202007%20Gene%20Pool%20Diffewrences%20between%20N%20S%20Alt.pdf |volume=43 |issue=5 |doi=10.1134/S1022795407050110 |pmid=17633562 |access-date=August 29, 2009 |archive-date=October 6, 2011 |archive-url=https://web.archive.org/web/20111006230239/http://www.medgenetics.ru/UserFile/File/Doc/Evolution%20Doc/Kharkov%202007%20Gene%20Pool%20Diffewrences%20between%20N%20S%20Alt.pdf |url-status=live }}
• {{Cite journal |vauthors=King RJ, Ozcan SS, Carter T, Kalfoğlu E, Atasoy S, Triantaphyllidis C, Kouvatsi A, Lin AA, Chow CE, Zhivotovsky LA, Michalodimitrakis M, Underhill PA |title=Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic |journal=Annals of Human Genetics |volume=72 |issue=Pt 2 |pages=205–214 |year=2008 |url=http://dirkschweitzer.net/E3b-papers/KingAHG-08-72-205.pdf |doi=10.1111/j.1469-1809.2007.00414.x |pmid=18269686 |s2cid=22406638 |display-authors=8 |url-status=dead |archive-url=https://web.archive.org/web/20090305052142/http://dirkschweitzer.net/E3b-papers/KingAHG-08-72-205.pdf |archive-date=March 5, 2009 }}
• {{Cite journal |doi=10.1098/rspb.2004.2698 |title=Unravelling migrations in the steppe: mitochondrial DNA sequences from ancient central Asians |first7=J. |last7=Bertranpetit |first6=D. |last6=Pettener |first5=G |last5=Besio |first4=P |last4=Mainardi |first3=C |last3=Mao |first2=M |last2=Robello |first1=C. |last1=Lalueza-Fox |journal=Proc. Biol. Sci. |year=2004 |volume=271 |pmid=15255049 |issue=1542 |pages=941–947 |pmc=1691686}}
• {{Cite journal |last1=Luca |first1=F |last2=Di Giacomo |first2=F |last3=Benincasa |first3=T |last4=Popa |first4=LO |last5=Banyko |first5=J |last6=Kracmarova |first6=A |last7=Malaspina |first7=P |last8=Novelletto |first8=A |last9=Brdicka |first9=R |s2cid=29801640 |title=Y-Chromosomal Variation in the Czech Republic |journal=American Journal of Physical Anthropology |volume=132 |issue=1 |year=2006 |pmid=17078035 |doi=10.1002/ajpa.20500 |pages=132–9|hdl=2108/35058 |hdl-access=free }}
• {{Cite journal |last1=Marjanovic |first1=D |last2=Fornarino |first2=S |last3=Montagna |first3=S |last4=Primorac |first4=D. |last5=Hadziselimovic |first5=R. |last6=Vidovic |first6=S. |last7=Pojskic |first7=N. |last8=Battaglia |first8=V. |last9=Achilli |first9=A. |last10=Drobnic |first10=K. |last11=Andjelinovic |first11=S. |last12=Torroni |first12=A. |last13=Santachiara-Benerecetti |first13=A. S. |last14=Semino |first14=O. |title=The peopling of modern Bosnia-Herzegovina: Y-chromosome haplogroups in the three main ethnic groups |journal=Annals of Human Genetics |volume=69 |issue=Pt 6 |pages=757–63 |date=November 2005 |pmid=16266413 |doi=10.1111/j.1529-8817.2005.00190.x |s2cid=36632274 |url=http://www3.interscience.wiley.com/resolve/openurl?genre=article&sid=nlm:pubmed&issn=0003-4800&date=2005&volume=69&issue=Pt%206&spage=757 |display-authors=8}}{{dead link|date=January 2025|bot=medic}}{{cbignore|bot=medic}}
• {{Cite journal |last1=Mukherjee |first1=Namita |publication-date=December 2001 |last2=Nebel |year=2001 |first2=Almut |last3=Oppenheim |first3=Ariella |last4=Majumder |first4=Partha P. |title=High-resolution analysis of Y-chromosomal polymorphisms reveals signatures of population movements from central Asia and West Asia into India |periodical=Journal of Genetics |volume=80 |issue=3 |pages=125–135 |doi=10.1007/BF02717908 |pmid=11988631 |s2cid=13267463 |url=http://www.ias.ac.in/jgenet/Vol80No3/125.pdf |access-date=September 2, 2019 |archive-date=September 2, 2019 |archive-url=https://web.archive.org/web/20190902163435/https://www.ias.ac.in/jgenet/Vol80No3/125.pdf |url-status=live }}
• {{Cite journal |doi=10.1086/324070 |first1=Almut |last1=Nebel |first2=Dvora |last2=Filon |first3=Bernd |last3=Brinkmann |first4=Partha |last4=Majumder |first5=Marina |last5=Faerman |first6=Ariella |last6=Oppenheim |title=The Y Chromosome Pool of Jews as Part of the Genetic Landscape of the Middle East |journal=American Journal of Human Genetics |year=2001 |volume=69 |issue=5 |pages=1095–112 |pmid=11573163 |pmc=1274378}}
• {{Cite journal |last1=Passarino |first1=G |last2=Semino |last3=Magria |last4=Al-Zahery |year=2001 |title=The 49a,f haplotype 11 is a new marker of the EU19 lineage that traces migrations from northern regions of the black sea |journal=Hum. Immunol. |volume=62 |issue=9 |pages=922–932 |doi=10.1016/S0198-8859(01)00291-9 |pmid=11543894 |first2=Ornella |first3=Chiara |first4=Nadia |last5=Benuzzi |first5=Giorgia |last6=Quintana-Murci |first6=Lluis |last7=Andellnovic |first7=Slmun |last8=Bullc-Jakus |first8=Floriana |last9=Liu |first9=Aiping |last10=Arslan |first10=Ahmet |last11=Santachiara-Benerecetti |first11=A. Silvana |display-authors=8}}
• {{Cite journal |last1=Qamar |year=2002 |first1=R |issue=5 |last2=Ayub |volume=70 |journal=American Journal of Human Genetics |first2=Q |last3=Mohyuddin |first3=A |last4=Helgason |first4=A |last5=Mazhar |first5=K |last6=Mansoor |first6=A |last7=Zerjal |first7=T |last8=Tylersmith |first8=C |last9=Mehdi |first9=S |title=Y-Chromosomal DNA Variation in Pakistan |doi=10.1086/339929 |pmid=11898125 |pmc=447589 |pages=1107–24}}
• {{Cite journal |doi=10.1086/318200 |last1=Quintana-Murci |first1=L |year=2001 |last2=Krausz |first2=C |last3=Zerjal |first3=T |last4=Sayar |first4=SH |last5=Hammer |first5=MF |last6=Mehdi |first6=SQ |last7=Ayub |first7=Q |last8=Qamar |first8=R |last9=Mohyuddin |first9=A |last10=Radhakrishna |first10=Uppala |last11=Jobling |first11=Mark A. |last12=Tyler-Smith |first12=Chris |last13=McElreavey |first13=Ken |title=Y-chromosome lineages trace diffusion of people and languages in southwestern Asia |journal=American Journal of Human Genetics |volume=68 |issue=2 |pages=537–542 |pmid=11133362 |pmc=1235289 |display-authors=8}}
• {{Cite journal |last1=Rebala |first1=Krzysztof |last2=Mikulich |first2=AI |last3=Tsybovsky |first3=IS |last4=Siváková |first4=D |last5=Džupinková |first5=Z |last6=Szczerkowska-Dobosz |first6=A |last7=Szczerkowska |first7=Z |title=Y-STR variation among Slavs: evidence for the Slavic homeland in the middle Dnieper basin |journal=Journal of Human Genetics |volume=52 |issue=5 |year=2007 |pmid=17364156 |doi=10.1007/s10038-007-0125-6 |pages=406–414 |doi-access=free }}
• {{Cite journal |last1=Saha |first1=Anjana |last2=Sharma |year=2005 |first2=S |last3=Bhat |first3=A |last4=Pandit |first4=A |last5=Bamezai |first5=R |title=Genetic affinity among five different population groups in India reflecting a Y-chromosome gene flow |periodical=Journal of Human Genetics |volume=50 |issue=1 |pages=49–51 |pmid=15611834 |doi=10.1007/s10038-004-0219-3|doi-access=free }}
• {{Cite journal |last1=Sanchez |year=2003 |first1=J |last2=Børsting |first2=C |last3=Hallenberg |first3=C |last4=Buchard |first4=A |last5=Hernandez |first5=A |last6=Morling |first6=N |title=Multiplex PCR and minisequencing of SNPs—a model with 35 Y chromosome SNPs |journal=Forensic Sci Int |pmid=14550618 |volume=137 |issue=1 |pages=74–84 |doi=10.1016/S0379-0738(03)00299-8}}
• {{Cite journal |last1=Soares |year=2010 |first1=Pedro |journal=Current Biology |pages=R174–83 |issue=4 |last2=Achilli |volume=20 |first2=Alessandro |doi=10.1016/j.cub.2009.11.054 |last3=Semino |first3=Ornella |last4=Davies |first4=William |last5=MacAulay |first5=Vincent |last6=Bandelt |first6=Hans-JüRgen |last7=Torroni |first7=Antonio |last8=Richards |first8=Martin B. |s2cid=7679921 |title=The Archaeogenetics of Europe |pmid=20178764 |doi-access=free |bibcode=2010CBio...20.R174S }}
• {{Cite journal |title=Genetic affinities among the lower castes and tribal groups of India: inference from Y chromosome and mitochondrial DNA |first1=Ismail |last1=Thanseem |first2=Kumarasamy |last2=Thangaraj |first3=Gyaneshwer |last3=Chaubey |first4=Vijay Kumar |last4=Singh |first5=Lakkakula VKS |last5=Bhaskar |first6=B Mohan |last6=Reddy |first7=Alla G |last7=Reddy |first8=Lalji |last8=Singh |journal=BMC Genetics |year=2006 |volume=7 |doi=10.1186/1471-2156-7-42 |pmid=16893451 |pmc=1569435 |page=42 |doi-access=free }}
• {{Cite journal |last1=Völgyi |first1=Antónia |last2=Zalán |year=2008 |first2=Andrea |last3=Szvetnik |first3=Enikő |last4=Pamjav |first4=Horolma |title=Hungarian population data for 11 Y-STR and 49 Y-SNP markers |pages=e27–8 |issue=2 |journal=Forensic Science International: Genetics |volume=3 |doi=10.1016/j.fsigen.2008.04.006 |pmid=19215861}}
• {{Cite journal |last1=Weale |year=2001 |first1=Michael |last2=Yepiskoposyan |first2=L |last3=Jager |first3=RF |last4=Hovhannisyan |first4=N |last5=Khudoyan |first5=A |last6=Burbage-Hall |first6=O |last7=Bradman |first7=N |last8=Thomas |first8=MG |s2cid=23113666 |title=Armenian Y chromosome haplotypes reveal strong regional structure within a single ethno-national group |journal=Hum Genet |pmid=11810279 |volume=109 |issue=6 |pages=659–674 |doi=10.1007/s00439-001-0627-9 |url=http://dirkschweitzer.net/E3b-papers/Weale-HG-01-659-Armenia-y-haplogroups.pdf }}{{dead link|date=October 2017 |bot=InternetArchiveBot |fix-attempted=yes }}
• {{Cite journal |last1=Weale |year=2002 |first1=S |last2=Zhivotovsky |first2=LA |last3=King |first3=R |last4=Mehdi |first4=SQ |last5=Edmonds |first5=CA |last6=Chow |first6=CE |last7=Lin |first7=AA |last8=Mitra |first8=M |last9=Sil |first9=SK |title=Y Chromosome Evidence for Anglo-Saxon Mass Migration |periodical=Mol. Biol. Evol. |volume=19 |issue=7 |pages=1008–1021 |pmid=12082121 |doi=10.1093/oxfordjournals.molbev.a004160 |doi-access=free }}
• {{Cite journal |last1=Wells |year=2001 |first1=R. S. |last2=Yuldasheva |first2=N |last3=Ruzibakiev |first3=R |last4=Underhill |first4=PA |last5=Evseeva |first5=I |last6=Blue-Smith |first6=J |last7=Jin |first7=L |last8=Su |first8=B |last9=Pitchappan |first9=R |last10=Shanmugalakshmi |first10=S. |last11=Balakrishnan |first11=K. |last12=Read |first12=M. |last13=Pearson |first13=N. M. |last14=Zerjal |first14=T. |last15=Webster |first15=M. T. |last16=Zholoshvili |first16=I. |last17=Jamarjashvili |first17=E. |last18=Gambarov |first18=S. |last19=Nikbin |first19=B. |last20=Dostiev |first20=A. |last21=Aknazarov |first21=O. |last22=Zalloua |first22=P. |last23=Tsoy |first23=I. |last24=Kitaev |first24=M. |last25=Mirrakhimov |first25=M. |last26=Chariev |first26=A. |last27=Bodmer |first27=W. F. |title=The Eurasian Heartland: A continental perspective on Y-chromosome diversity |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=98 |issue=18 |pages=10244–9 |pmid=11526236 |pmc=56946 |doi=10.1073/pnas.171305098 |bibcode=2001PNAS...9810244W |display-authors=8|doi-access=free }}
• {{Cite journal |last1=Wilson |first1=J. F. |last2=Weiss |year=2001 |first2=DA |last3=Richards |first3=M |last4=Thomas |first4=MG |last5=Bradman |first5=N |last6=Goldstein |first6=DB |title=Genetic evidence for different male and female roles during cultural transitions in the British Isles |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=98 |pages=5078–5083 |doi=10.1073/pnas.071036898 |issue=9 |pmc=33166 |pmid=11287634 |bibcode=2001PNAS...98.5078W|doi-access=free }}
• {{Cite journal |last1=Zerjal |year=2001 |first1=T |last2=Beckman |first2=L |last3=Beckman |first3=G |last4=Mikelsaar |first4=AV |last5=Krumina |first5=A |last6=Kucinskas |first6=V |last7=Hurles |first7=ME |last8=Tyler-Smith |first8=C |title=Geographical, linguistic, and cultural influences on genetic diversity: Y-chromosomal distribution in Northern European populations |journal=Mol Biol Evol |volume=18 |issue=6 |pages=1077–1087 |pmid=11371596 |doi=10.1093/oxfordjournals.molbev.a003879|doi-access=free }}
• {{Cite journal |last1=Zerjal |first1=T |last2=Wells |first2=RS |last3=Yuldasheva |first3=N |last4=Ruzibakiev |first4=R |last5=Tyler-Smith |first5=C |title=A Genetic Landscape Reshaped by Recent Events: Y-Chromosomal Insights into Central Asia |journal=American Journal of Human Genetics |volume=71 |issue=3 |pages=466–482 |year=2002 |pmid=12145751 |pmc=419996 |doi=10.1086/342096 }}
• {{Cite journal |last1=Zhao |first1=Zhongming |last2=Khan |first2=Faisal |last3=Borkar |first3=Minal |last4=Herrera |first4=Rene |last5=Agrawal |first5=Suraksha |title=Presence of three different paternal lineages among North Indians: A study of 560 Y chromosomes |journal=Annals of Human Biology |year=2009 |pages=1–14 |issue=1 |doi=10.1080/03014460802558522 |volume=36 |pmid=19058044 |pmc=2755252}}
• {{Cite journal |doi=10.1086/380911 |last1=Zhivotovsky |first1=L |last2=Underhill |first2=PA |last3=Cinnioğlu |first3=C |last4=Kayser |first4=M |last5=Morar |first5=B |last6=Kivisild |first6=T |last7=Scozzari |first7=R |last8=Cruciani |first8=F |last9=Destro-Bisol |first9=G |last10=Spedini |first10=G |last11=Chambers |first11=G. K. |last12=Herrera |first12=R. J. |last13=Yong |first13=K. K. |last14=Gresham |first14=D |last15=Tournev |first15=I |last16=Feldman |first16=M. W. |last17=Kalaydjieva |first17=L |title=The effective mutation rate at Y chromosome short tandem repeats, with application to human population-divergence time |journal=American Journal of Human Genetics |year=2004 |volume=74 |issue=1 |pages=50–61 |pmid=14691732 |pmc=1181912 |display-authors=8}}
• {{cite journal|author=G. Mršić|title=Croatian national reference Y-STR haplotype database |journal=Molecular Biology Reports|volume=39|issue=7|date=2012|doi=10.1007/s11033-012-1610-3|pmid=22391654|ref={{harvid|Mršić et al.|2012}}|display-authors=etal|pages=7727–41|s2cid=18011987}}
• {{cite journal|author=J. Šarac|title=Genetic heritage of Croatians in the Southeastern European gene pool—Y chromosome analysis of the Croatian continental and Island population|journal= American Journal of Human Biology|volume=28|issue=6|date=2016|doi=10.1002/ajhb.22876|pmid=27279290|ref={{harvid|Šarac et al.|2016}}|display-authors=etal|pages=837–845|s2cid=25873634}}

{{Refend}}

{{collapse bottom}}

{{Commons category|Haplogroup R1a of Y-DNA}}

;DNA Tree

• [http://www.familytreedna.com/public/R1aY%2DHaplogroup/ FTDNA R1a Y-chromosome Haplogroup Project]
• [http://www.familytreedna.com/public/R1a/default.aspx R1a1a1 and Subclades Y-DNA Project – Background] Family Tree DNA R1a1a1

;TMRCA

• [http://taylorfamilygenes.info/TMRCA.shtml TMRCA = Time to Most Recent Common Ancestor]

;Various

• [http://danishdemes.org/YDNA-results-HgR1a.shtml Danish Demes Regional DNA Project: Y-DNA Haplogroup R1a] {{Webarchive|url=https://web.archive.org/web/20080820071422/http://danishdemes.org/YDNA-results-HgR1a.shtml |date=August 20, 2008 }}
• Eurogenes Blog, [http://eurogenes.blogspot.nl/2016/01/the-poltavka-outlier.html The Poltovka outlier]
• Avotaynu Online, [http://www.avotaynuonline.com/2016/07/identifying-the-genetic-fingerprint-of-a-tzaddik-that-touched-the-world-the-shpoler-zeida/ The Y-DNA Fingerprint of the Shpoler Zeida, a Tzaddik Who Touched the World]

{{DEFAULTSORT:Haplogroup R1a (Y-Dna)}}

M420

R