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Haplogroup D-M174

From Wikipedia, the free encyclopedia
Human Y-chromosome DNA haplogroup

Haplogroup D-M174
Possible time of origin50,000[1]-60,000[2] years BP

65,200 [95% CI 62,100 <-> 68,300] ybp[3]
Coalescence age46,300 [95% CI 43,500 <-> 49,100] ybp[3]
Possible place of originAsia[2][4][5] (probablySouth Asia[6] orCentral Asia[7])
AncestorD (CTS3946)
DescendantsD1a (CTS11577)
D1a1 (Z27276)
D1a2 (Z3660)
D1a2a (M55)
D1a2b (Y34637)
D1b (L1378)
Defining mutationsM174, IMS-JST021355, PAGES00003

Haplogroup D1 orD-M174 is asubclade ofhaplogroup D-CTS3946. This broad lineage of the male-specific portion of the human Y chromosome is found primarily inEast Asia,Magar-ethnicNepal and theAndaman Islands. It is also found regularly with lower frequency inCentral Asia,North Asia andMainland Southeast Asia, and, more rarely, inEurope andWest Asia.

Origins

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Human Y chromosome haplogroup D-M174 likely originated inAsia roughly 60,000 years ago.[2][4] While haplogroup D-M174, along withhaplogroup E, contains the distinctiveYAPpolymorphism—which indicates their closer ancestry than C—no haplogroup D-M174chromosomes have been found outside of Asia.[4] Haplogroup D1 is also often found in Southern Asia's populations.[8]

Several studies (Hammer et al. 2006, Shinoda 2008, Matsumoto 2009) suggested that paternal haplogroup D-M174 originated inCentral Asia.[7]

A 2017 study by Mondal et al. finds that theRiang people (aTibeto-Burmese population) and theAndamanese share the same D clade (D1a3, also known as D1a2b) and have their closest lineages with other clades in East Asia. TheJarawa andOnge shared D1a2b with each other within the last ~7,000 years. The East Asian D1a2b diverged from theJapanese D1a2a lineage ~53,000 years ago. The authors conclude: "This strongly suggests that haplogroup D does not indicate a separate ancestry for Andamanese populations. Rather, haplogroup D was part of the standing variation carried by the Eastern OOA expansion, and later lost from most of the populations except in Andaman and partially in Japan and Tibet".[9]

A 2019 study by Haber et al. suggested that Haplogroup D-M174 originated in Central Asia and evolved as it migrated to different directions of the continent. One group of population migrated to Siberia, others to Japan and Tibet, and another group migrated to the Andaman islands.[10]

A 2020 genetic study by Hallast et al. on ancient and modern haplogroups using aphylogenetic analysis of haplogroup C, D, and FT sequences—including very rare deep-rooting lineages such as D0/D2, a divergent D lineage not belonging to D-M174—argues that the initial splits withinhaplogroup CT (an ancestor ofDE) occurred inAfrica. It also argues thatphylogeographic analyses of ancient and present-day non-AfricanY chromosomes all point to East/Southeast Asia as the origin of all known surviving non-African male lineages (apart from recent migrants) soon after an initial 70,000–55,000-year-ago migration from Africa ofbasal haplogroup D and other basal Y-lineages. It argues that these lineages then rapidly expanded acrossEurasia, diversified in Southeast Asia, and expanded westward around 55,000–50,000 years ago, replacing other local lineages within Eurasia; haplogroup D (as D-M174) then underwent rapid expansions within Eastern Eurasian populations and consists of five branches that formed about 45,000 years ago. The study finds that these haplogroups currently have their greatest diversity in Eastern Eurasia (East/Southeast Asia). Tibeto-Burmese populations of East and Southeast Asia were found to have the highest amount of diversity.[11]

A 2024 study found that a primitive D1b subclade can be found in the Philippines and possibly in MalaysianHoabinhian foragers.[12]

Overview

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Haplogroup D-M174 is found today with high frequency among populations inTibet,Magar-ethnicNepal, northernMyanmar,Qinghai, theJapanese archipelago, and theAndaman Islands, though curiously not as much in the rest ofIndia. TheAinu people ofJapan and various Tibeto-Burmese people (such as theTripuri people) are notable for possessing almost exclusively haplogroup D-M174 chromosomes. Haplogroup D-M174 chromosomes are also found at low to moderate frequencies among theBai,Dai,Han,Hui,Manchu,Miao,Tujia,Xibe,Yao, andZhuang peoples ofChina and among several minority populations ofSichuan andYunnan that speakTibeto-Burman languages and reside in close proximity to the Tibetans, such as theJingpo,Jino,Mosuo,Naxi,Pumi,Qiang, andYi.[13]

Haplogroup D is also found in populations in China proper and inKorea, but with much lower frequency than in Tibet and Japan. A study published in 2011 found D-M174 in 2.49% (43/1729) of Han Chinese males, with frequencies tending to be higher than average toward the north and west of the country (8.9% ofShaanxi Han, 5.9% ofGansu Han, 4.4% ofYunnan Han, 3.7% ofGuangxi Han, 3.3% ofHunan Han, and 3.2% ofSichuan Han).[14] In another study of Han Chinese Y-DNA published in 2011, haplogroup D-M174 was observed in 1.94% (7/361) of a sample of unrelated Han Chinese male volunteers atFudan University inShanghai, with the origins of most of the volunteers being traced back to East China (Jiangsu,Zhejiang, Shanghai, andAnhui).[15]

In Korea, haplogroup D-M174 was observed in 3.8% (5/133) of a sample fromDaejeon,[16] 3.5% (3/85) of a sample fromSeoul,[17] 3.3% (3/90) of a sample fromJeolla,[18] 2.4% (2/84) of a sample fromGyeongsang,[18] 2.3% (13/573) of another sample from Seoul,[16] 1.4% (1/72) of a sample fromChungcheong,[18] 1.1% (1/87) of a sample fromJeju,[18] and 0.9% (1/110) of a third sample from Seoul-Gyeonggi.[18] In other studies, haplogroup D-M174 has been observed in 6.7% (3/45)[19] and 4.0% (3/75)[20] of samples from Korea without any further specification of the area of sampling.

Little high-resolution data regarding the phylogenetic position of Han Chinese and Korean members of Y-DNA haplogroup D has been published, but the available data suggests that most Han Chinese members of haplogroup D should belong to clades found frequently among Tibetans (especially the D-M15 clade, also found among speakers of someLolo-Burmese andHmong-Mien languages), whereas most Korean members of haplogroup D should belong to the D-M55 clade, which is found frequently amongAinu,Ryukyuan, andJapanese people.[20][18][3]

Haplogroup D Y-DNA has been found (albeit with low frequency) among modern populations of theEurasian steppe, such as:

It has also been found among linguistically similar (Turkic- orMongolic-speaking) modern populations of the desert and oasis belt south of the steppe, such asYugurs,Bao'an,Monguors,Uyghurs, andUzbeks. In commercial testing, members have been found as far west asRomania in Europe andIraq in Western Asia.[26]

Unlike haplogroup C-M217, haplogroup D-M174 is not found in theNew World; it is not present in any modern Native American (North, Central, or South) populations. While it is possible that it traveled to the New World like haplogroup C-M217, those lineages apparently became extinct.

Haplogroup D-M174 is remarkable for its rather extreme geographic differentiation, with a distinct subset of chromosomes being found exclusively in each of the populations that contains a large percentage of individuals whose Y-chromosomes belong to haplogroup D-M174:haplogroup D-M15 amongTibetans, as well as other East/Southeast Asian populations that display low frequencies of haplogroup D-M174 Y-chromosomes;haplogroup D-M55 among the various populations of the Japanese archipelago and with low frequency amongKoreans; and haplogroup D-P99 among the inhabitants of Tibet and some other parts of central Eurasia (e.g. Mongolia[27] and theAltai[20][21][22]). D-M174* without positive-tested subclades D-M15 or D-M55 is found at high frequencies amongAndaman Islanders, and recently an Andamanese subclade was found to beD-Y34637 (D1a2b).[3] Another type (or types) of paragroup D-M174* without positive-tested subclades of D-M15, D-P47, or D-M55 is found at a very low frequency among theTurkic andMongolic populations ofCentral Asia, amounting to no more than 1% in total. This apparently ancient diversification of haplogroup D-M174 suggests that it may perhaps be better characterized as a "super-haplogroup" or "macro-haplogroup".

In one study, the frequency of haplogroup D-M174 without positive-tested subclades found amongThais was 10%.[2] Suet al. (2000) found DE-YAP/DYS287(xM15) in 11.1% (5/45) of a set of three samples from Thailand—including 20% (4/20) North Thai, 20% (1/5)So, and 0% (0/20) Northeast Thai—and in 16.7% (1/6) of a sample fromGuam.[28] Meanwhile, the authors found D-M15 in 15% of a pair of samples ofYao, including 30% (3/10) YaoJinxiu and 0% (0/10) YaoNandan; 14.3% (2/14) of a sample ofYi; 3.8% (1/26) of a sample ofCambodians; and 3.6% (1/28) of a sample ofZhuang.[28] Donget al. (2002) found DE-YAP Y-chromosomes in 12.5% (2/16) of a sample ofJingpo fromLuxi City, Yunnan, 10.0% (2/20) of a sample ofDai from Luxi City, and 1.82% (1/55) of a sample ofNu fromGongshan andFugong, Yunnan.[29]

Distribution and subclades

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The haplogroup D-M174 Y-chromosomes that are found among Tibeto-Burman populations as well as people of the Japanese archipelago belong to haplogroup D1a2b, D1a2a, and D1a1. D-M55 (D1a2a) is particularly distinctive, bearing a complex of at least five individual mutations along an internal branch of the haplogroup D-M174phylogeny, thus distinguishing it clearly from the other haplogroup D-M174 chromosomes that are found among Tibetans and Andaman Islanders and providing evidence that Y-chromosome haplogroup D-M55 was the modal haplogroup in the ancestral population that developed the prehistoricJōmon culture in the Japanese islands.

It is suggested that the majority of D-M174 Y-chromosome carriers migrated fromCentral Asia toEast Asia. One group migrated to the Andaman Islands, thus forming or helping to form the Andamanese people. Another group stayed in modern Tibet and southern China (today Tibeto-Burman peoples), and a third group migrated to Japan, possibly via theKorean Peninsula (pre-Jōmon people).[2][20]

D-Z27276 (D1a1)

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Haplogroup D-Z27276 is the common ancestor of D-M15 and D-P99, which are common in Tibet (China).

D-M15 (D1a1a)

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D-M15 was first reported to have been found in a sample fromCambodia andLaos (1/18 = 5.6%) and in a sample from Japan (1/23 = 4.3%) in a preliminary worldwide survey of Y-DNA variation in extant human populations.[30]

Subsequently, Y-DNA belonging to haplogroup D-M15 has been found frequently amongTibeto-Burman-speaking populations ofSouthwestern China (including approximately 23% ofQiang,[2][31][32] approximately 12.5% ofTibetans,[2] and approximately 9% ofYi[2][33]), and amongYao people inhabiting northeasternGuangxi (6/31 = 19.4% Lowland Yao, 5/41 = 12.2% Native Mien, 3/41 = 7.3% Lowland Kimmun),[34] with a moderate distribution throughoutCentral Asia,East Asia, and continentalSoutheast Asia (Indochina).[2]

A study published in 2011 found D-M15 in 7.8% (4/51) of a sample ofHmong Daw and in 3.4% (1/29) of a sample ofXinhmul from northern Laos.[34]

D-P47 (D1a1b1)

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This subclade is found with high frequency amongPumi,[2]Naxi,[2] andTibetans,Lu D, Lou H, Yuan K, Wang X, Wang Y, Zhang C, et al. (September 2016)."Ancestral Origins and Genetic History of Tibetan Highlanders".American Journal of Human Genetics.99 (3):580–594.doi:10.1016/j.ajhg.2016.07.002.PMC 5011065.PMID 27569548.[2] with a moderate distribution inCentral Asia.[2] According to one study, Tibetans have a frequency of about 41.31% of haplogroup D-P47.[2]

D-Z3660 (D1a2)

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For about 7,000 years, the natives of the Andaman Islands shared a common ancestry with each other. The closest lineage to the Andamanese is the Japanese haplogroup D, with which it has a very old relationship, dating back to about 53,000 years.[9]

D-M55 (D1a2a)

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Previously known asD-M55, D-M64.1/Page44.1 (D1a2a) is found with high frequency amongAinu[35] and with medium frequency amongJapanese[36] andRyukyuans.[36]

Kimet al. (2011) found haplogroup D-M55 in 2.0% (1/51) of a sample ofBeijingHan and in 1.6% (8/506) of a pool of samples fromSouth Korea, including 3.3% (3/90) from the Jeolla region, 2.4% (2/84) from the Gyeongsang region, 1.4% (1/72) from the Chungcheong region, 1.1% (1/87) from the Jeju region, 0.9% (1/110) from the Seoul-Gyeonggi region, and 0% (0/63) from the Gangwon region.[18] Hammeret al. (2006) found haplogroup D-P37.1 in 4.0% (3/75) of a sample from South Korea.[20]

D-M116.1, which is a subclade of D-M55, has been observed in one individual in a sample from Micronesia (n=17) according to the supplementary material of a study published in 2006.[20] D-M116.1 also has been observed in one individual in a pool of samples from West Timor (n=497); the pertinent individual is from Umaklaran, located on the north side of the island of Timor near the border withEast Timor.[37]

According toMitsuru Sakitani, haplogroup D1 arrived from Central Asia to northernKyushu via theAltai Mountains and theKorean Peninsula more than 40,000 years ago, and haplogroup D-M55 (D1a2a) was born in the Japanese archipelago.[38]

D-Y34637 (D1a2b)

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D1a2b (formerly one of D*) is found at high frequencies amongAndaman Islanders,[2] especiallyOnge (23/23 = 100%) andJarawa (4/4 = 100%).[39][3]

D-L1378 (D1b)

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D1b (L1378, M226.2) has been found in commercial testing in two families fromMactan Island in theCebu region of thePhilippines, in the ethnicRade people fromVietnam as well as an ancient sample from Malaysia.[40]

D-M174*

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D-M174 (xM15, P99, M55) is found in some tribes ofNortheast India (among whom rates vary from 0% to 65%).[5][41][42][43]

The basal D-M174 (xM15, P47, M55) has been found in approximately 5% ofAltaians.[20] Kharkovet al. found haplogroup D*(xD-M15) in 6.3% (6/96) of a pool of samples of Southern Altaians from three different localities, particularly inKulada (5/46 = 10.9%) andKosh-Agach (1/7 = 14%), though they did not test for any marker of the subclade D-M55 or D-P99. Kharkovet al. also reported finding haplogroup DE-M1(xD-M174) Y-DNA in one Southern Altaian individual fromBeshpeltir (1/43 = 2.3%).[21]

In 2023, D-M174 was found in one individual inLa Crosse, Wisconsin.[44]

Phylogenetics

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Phylogenetic history

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Main article:Conversion table for Y chromosome haplogroups

Prior to 2002, there were in academic literature at least seven naming systems for the Y-chromosome phylogenetic tree. This led to considerable confusion. In 2002, major research groups came together and formed theY Chromosome Consortium (YCC). They published a joint paper that created a single new tree. Later, a group of citizen scientists with an interest in population genetics and genealogy formed a working group to create an amateur tree. The table below brings together all of these works at the point of the landmark 2002 YCC tree. This allows a researcher reviewing older published literature to quickly move between nomenclatures.

YCC 2002/2008 (Shorthand)(α)(β)(γ)(δ)(ε)(ζ)(η)YCC 2002 (Longhand)YCC 2005 (Longhand)YCC 2008 (Longhand)YCC 2010r (Longhand)ISOGG 2006ISOGG 2007ISOGG 2008ISOGG 2009ISOGG 2010ISOGG 2011ISOGG 2012
D-M174********DDDDDDDDDD
D-M154IV3G12Eu5H3BD1D1D1D1D1D1D1D1D1D1D1
D-M55********D2D2D2D2D2D2D2D2D2D2
D-P124IV3G11Eu5H2BD2aD2aD2a1a1D2a1a1D2D2D2a1a1D2a1a1D2a1a1removedremoved
D-M116.14IV3G11Eu5H2BD2b*D2aD2aD2aD2aD2aD2aD2aD2aremovedremoved
D-M1254IV3G11Eu5H2BD2b1D2a1D2a1D2a1D2a1D2a1D2a1D2a1D2a1D2a1D2a1
D-M1514IV3G11Eu5H2BD2b2D2a1D2a2D2a2D2a2D2a2D2a2D2a2D2a2D2a2D2a2

Research publications

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The following research teams, per their publications, were represented in the creation of the YCC tree.

  • α Jobling and Tyler-Smith (2000)[45] and Kalaydjieva (2001)[46]
  • β Underhill (2000)[47]
  • γ Hammer (2001)[48]
  • δ Karafet (2001)[49]
  • ε Semino (2000)[50]
  • ζ Su (1999)[51]
  • η Capelli (2001)[52]

Phylogenetic trees

[edit]

By ISOGG tree(Version 14.151):[53]


See also

[edit]

Genetics

[edit]

Y-DNA D-M174 subclades

[edit]

Y-DNA backbone tree

[edit]
This article needs to beupdated. Please help update this article to reflect recent events or newly available information.(February 2021)
Footnotes
  1. ^Van Oven M, Van Geystelen A, Kayser M, Decorte R, Larmuseau HD (2014). "Seeing the wood for the trees: a minimal reference phylogeny for the human Y chromosome".Human Mutation.35 (2):187–91.doi:10.1002/humu.22468.PMID 24166809.S2CID 23291764.
  2. ^International Society of Genetic Genealogy (ISOGG; 2015),Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.)
  3. ^Haplogroup A0-T is also known as A-L1085 (and previously as A0'1'2'3'4).
  4. ^Haplogroup A1 is also known as A1'2'3'4.
  5. ^ F-Y27277, sometimes known as F2'4, is both the parent clade of F2 and F4 and a child of F-M89.
  6. ^Haplogroup LT (L298/P326) is also known as Haplogroup K1.
  7. ^Between 2002 and 2008,Haplogroup T-M184 was known as "Haplogroup K2". That name has since been re-assigned toK-M526, the sibling of Haplogroup LT.
  8. ^ Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.
  9. ^ Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure.
  10. ^ Haplogroup P (P295) is also klnown as K2b2.
  11. ^K-M2313*, which as yet has no phylogenetic name, has been documented in two living individuals, who have ethnic ties to India and South East Asia. In addition, K-Y28299, which appears to be a primary branch of K-M2313, has been found in three living individuals from India. See: Poznikop. cit.;YFull YTree v5.08, 2017, "K-M2335", and;PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (Access date of these pages: 9 December 2017)
  12. ^ Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)
  13. ^ Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)

References

[edit]
  1. ^"Y-DNA Haplogroup D-M174 and its Subclades - 2017".
  2. ^abcdefghijklmnoShi H, Zhong H, Peng Y, Dong YL, Qi XB, Zhang F, et al. (October 2008)."Y chromosome evidence of earliest modern human settlement in East Asia and multiple origins of Tibetan and Japanese populations".BMC Biology.6 (1) 45.Bibcode:2008BMCB....6...45S.doi:10.1186/1741-7007-6-45.PMC 2605740.PMID 18959782.
  3. ^abcde"D YTree".YFull. Archived fromthe original on 2019-08-31. Retrieved2018-03-30.
  4. ^abcKarafet TM, Mendez FL, Meilerman MB, Underhill PA, Zegura SL, Hammer MF (May 2008)."New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree".Genome Research.18 (5):830–8.doi:10.1101/gr.7172008.PMC 2336805.PMID 18385274.
  5. ^abSu B, Xiao C, Deka R, Seielstad MT, Kangwanpong D, Xiao J, et al. (December 2000). "Y chromosome haplotypes reveal prehistorical migrations to the Himalayas".Human Genetics.107 (6):582–90.doi:10.1007/s004390000406.PMID 11153912.S2CID 36788262.
  6. ^Xu D, Li H (5 May 2017).Languages and Genes in Northwestern China and Adjacent Regions. Springer. p. 25.ISBN 978-981-10-4169-3. "For the origin of haplogroup D, Chandrasekar et al. (2007) suggested that the CT-M168 gave rise to the YAP insertion and D-M174 mutation in South Asia based on their findings of the YAP insertion in northeast Indian tribes and the D-M174 in South Asia and the D-M174 in Andaman islanders."
  7. ^abMatsumoto H (February 2009)."The origin of the Japanese race based on genetic markers of immunoglobulin G".Proceedings of the Japan Academy. Series B, Physical and Biological Sciences.85 (2):69–82.Bibcode:2009PJAB...85...69M.doi:10.2183/pjab.85.69.PMC 3524296.PMID 19212099.
  8. ^Guo Y, Xia Z, Cui W, Chen C, Jin X, Zhu B (May 2020)."Joint Genetic Analyses of Mitochondrial and Y-Chromosome Molecular Markers for a Population from Northwest China".Genes.11 (5): 564.doi:10.3390/genes11050564.PMC 7290686.PMID 32443545.
  9. ^abMondal M, Bergström A, Xue Y, Calafell F, Laayouni H, Casals F, et al. (May 2017). "Y-chromosomal sequences of diverse Indian populations and the ancestry of the Andamanese".Human Genetics.136 (5):499–510.doi:10.1007/s00439-017-1800-0.hdl:10230/34399.PMID 28444560.S2CID 3725426.In contrast, the Riang (Tibeto-Burman-speaking) and Andamanese have their nearest neighbour lineages in East Asia. The Jarawa and Onge shared haplogroup D lineages with each other within the last ~7000 years, but had diverged from Japanese haplogroup D Y-chromosomes ~53000 years ago, most likely by a split from a shared ancestral population.
  10. ^Haber M, Jones AL, Connell BA, Arciero E, Yang H, Thomas MG, et al. (1 August 2019)."A Rare Deep-Rooting D0 African Y-Chromosomal Haplogroup and Its Implications for the Expansion of Modern Humans Out of Africa".Genetics.212 (4):1421–1428.doi:10.1534/genetics.119.302368.PMC 6707464.PMID 31196864.
  11. ^Hallast P, Agdzhoyan A, Balanovsky O, Xue Y, Tyler-Smith C (February 2021)."A Southeast Asian origin for present-day non-African human Y chromosomes".Human Genetics.140 (2):299–307.doi:10.1007/s00439-020-02204-9.PMC 7864842.PMID 32666166.
  12. ^Cabrera VM (2024)."Early male and female footprints of modern humans across Eurasia and Australasia".Medical Research Archives.12 (8).bioRxiv 10.1101/2024.05.27.596017.doi:10.18103/mra.v12i8.5552.
  13. ^"Y染色体单倍群D在東亞的分布及其意義" [Distribution and significance of Y chromosome haplogroup D in East Asia](PDF) (in Chinese). Archived fromthe original(PDF) on 3 March 2016.
  14. ^Zhong H, Shi H, Qi XB, Duan ZY, Tan PP, Jin L, et al. (2011)."Extended Y Chromosome Investigation Suggests Postglacial Migrations of Modern Humans into East Asia via the Northern Route".Mol. Biol. Evol.28 (1):717–727.doi:10.1093/molbev/msq247.PMID 20837606.
  15. ^Yan S, Wang CC, Li H, Li SL, Jin L, et al. (The Genographic Consortium) (September 2011)."An updated tree of Y-chromosome Haplogroup O and revised phylogenetic positions of mutations P164 and PK4".European Journal of Human Genetics.19 (9):1013–5.doi:10.1038/ejhg.2011.64.PMC 3179364.PMID 21505448.
  16. ^abPark MJ, Lee HY, Yang WI, Shin KJ (July 2012). "Understanding the Y chromosome variation in Korea--relevance of combined haplogroup and haplotype analyses".International Journal of Legal Medicine.126 (4):589–99.doi:10.1007/s00414-012-0703-9.PMID 22569803.S2CID 27644576.
  17. ^abcdKatoh T, Munkhbat B, Tounai K, Mano S, Ando H, Oyungerel G, et al. (February 2005). "Genetic features of Mongolian ethnic groups revealed by Y-chromosomal analysis".Gene.346:63–70.doi:10.1016/j.gene.2004.10.023.PMID 15716011.
  18. ^abcdefgKim SH, Kim KC, Shin DJ, Jin HJ, Kwak KD, Han MS, et al. (April 2011)."High frequencies of Y-chromosome haplogroup O2b-SRY465 lineages in Korea: a genetic perspective on the peopling of Korea".Investigative Genetics.2 (1): 10.doi:10.1186/2041-2223-2-10.PMC 3087676.PMID 21463511.
  19. ^abcWells RS, Yuldasheva N, Ruzibakiev R, Underhill PA, Evseeva I, Blue-Smith J, et al. (August 2001)."The Eurasian heartland: a continental perspective on Y-chromosome diversity".Proceedings of the National Academy of Sciences of the United States of America.98 (18):10244–9.Bibcode:2001PNAS...9810244W.doi:10.1073/pnas.171305098.PMC 56946.PMID 11526236.
  20. ^abcdefghHammer MF, Karafet TM, Park H, Omoto K, Harihara S, Stoneking M, et al. (2006)."Dual origins of the Japanese: common ground for hunter-gatherer and farmer Y chromosomes".Journal of Human Genetics.51 (1):47–58.doi:10.1007/s10038-005-0322-0.PMID 16328082.
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