Human Y-chromosome DNA haplogroup

In human genetics, a human Y-chromosome DNA haplogroup is a haplogroup defined by mutations in the non-recombining portions of DNA from the Y chromosome (called Y-DNA). Many people within a haplogroup share a type of mutation called single-nucleotide polymorphisms (SNPs).[1]

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The human Y-chromosome accumulates roughly two mutations per generation.[2] Y-DNA haplogroups represent major branches of the Y-chromosome phylogenetic tree that share hundreds or even thousands of mutations unique to each haplogroup.

The Y-chromosomal most recent common ancestor (Y-MRCA, informally known as Y-chromosomal Adam) is the most recent common ancestor (MRCA) from whom all currently living humans are descended patrilineally. Y-chromosomal Adam is estimated to have lived roughly 236,000 years ago in Africa. By examining other bottlenecks most Eurasian men (men from populations outside of Africa) are descended from a man who lived 69,000 years ago. Other major bottlenecks occurred about 50,000 and 5,000 years ago and subsequently the ancestry of most Eurasian/non-African men can be traced back to four ancestors who lived 50,000 years ago.[3][4][5]

Naming convention
Schematic illustration of Y-DNA haplogroups naming convention. Haplogroups are defined through mutations (SNPs).

Y-DNA haplogroups are defined by the presence of a series of Y-DNA SNP markers. Subclades are defined by a terminal SNP, the SNP furthest down in the Y-chromosome phylogenetic tree.[6][7] The Y Chromosome Consortium (YCC) developed a system of naming major Y-DNA haplogroups with the capital letters A through T, with further subclades named using numbers and lower case letters (YCC longhand nomenclature). YCC shorthand nomenclature names Y-DNA haplogroups and their subclades with the first letter of the major Y-DNA haplogroup followed by a dash and the name of the defining terminal SNP.[8]

Y-DNA haplogroup nomenclature is changing over time to accommodate the increasing number of SNPs being discovered and tested, and the resulting expansion of the Y-chromosome phylogenetic tree. This change in nomenclature has resulted in inconsistent nomenclature being used in different sources.[1] This inconsistency, and increasingly cumbersome longhand nomenclature, has prompted a move towards using the simpler shorthand nomenclature. In September 2012, Family Tree DNA provided the following explanation of its changing Y-DNA haplogroup nomenclature to individual customers on their Y-DNA results pages (note that the haplogroup mentioned below relates to a specific individual):[9]

Long time customers of Family Tree DNA have seen the YCC-tree of Homo Sapiens evolve over the past several years as new SNPs have been discovered. Sometimes these new SNPs cause a substantial change in the "longhand" explanation of your terminal Haplogroup. Because of this confusion, we introduced a shorthand version a few years ago that lists the branch of the tree and your terminal SNP, i.e. J-L147, in lieu of J1c3d. Therefore, in the very near term, Family Tree DNA will discontinue showing the current "longhand" on the tree and we will focus all of our discussions around your terminal defining SNP.
This changes no science – it just provides an easier and less confusing way for us all to communicate.

— 

Bennett Greenspan, Family Tree DNA
Dr. Michael F. Hammer, University of Arizona

Phylogenetic structure
Phylogenetic tree of Y-DNA haplogroups [10]
 

Y-Chromosomal Adam

 
 
 

Haplogroup A

 
BT
 

Haplogroup B

 
CT
DE
 

Haplogroup D

 
 

Haplogroup E

 
 
CF
 

Haplogroup C

 
F
 

Haplogroup G

 
HIJK
 

Haplogroup H

 
IJK
IJ
 

Haplogroup I

 
 

Haplogroup J

 
 
K
LT
 

Haplogroup L

 
 

Haplogroup T

 
 
MNOPS
NO
 

Haplogroup N

 
 

Haplogroup O

 
 
MPS
MS
 

Haplogroup S

 
 

Haplogroup M

 
 
P
 

Haplogroup Q

 
 

Haplogroup R

 
 
 
 
 
 
 
 
 
 
 
 

Major Y-DNA haplogroups

Haplogroups A and B

Haplogroup A is the NRY (non-recombining Y) macrohaplogroup from which all modern paternal haplogroups descend. It is sparsely distributed in Africa, being concentrated among Khoisan populations in the southwest and Nilotic populations toward the northeast in the Nile Valley. BT is a subclade of haplogroup A, more precisely of the A1b clade (A2-T in Cruciani et al. 2011), as follows:

  • Haplogroup A
    • Haplogroup A00
    • Haplogroup A0 (formerly also A1b)
    • Haplogroup A1 (also A1a-T)
      • Haplogroup A1a (M31)
      • Haplogroup A1b (also A2-T; P108, V221)
        • Haplogroup A1b1a1 (also A2; M14)
        • Haplogroup A1b1b (also A3; M32)
        • Haplogroup BT (M91, M42, M94, M139, M299)
          • Haplogroup B (M60)
          • Haplogroup CT

Haplogroup CT (P143)

The defining mutations separating CT (all haplogroups except for A and B) are M168 and M294. The site of origin is likely in Africa. Its age has been estimated at approximately 88,000 years old,[11][12] and more recently at around 100,000[13] or 101,000 years old.[14]

Haplogroup C (M130)

Haplogroup D (CTS3946)

Haplogroup E (M96)
  • Haplogroup E (M40, M96) Found in Africa and parts of the Middle East and Europe
    • Haplogroup E1 (P147)
      • Haplogroup E1a (M33, M132) formerly E1
      • Haplogroup E1b (P177)
        • Haplogroup E1b1 (P2, DYS391p); formerly E3
          • Haplogroup E1b1a (V38)
            • Haplogroup E1b1a1 (M2) Found in Africa, especially among Niger–Congo-speaking populations.; formerly E3a
            • Haplogroup E1b1a2 (M329) Found in Africa, especially in Ethiopia among Omotic-speaking populations.; formerly E3*
          • Haplogroup E1b1b (M215)
    • Haplogroup E2 (M75)

Haplogroup F (M89)

The groups descending from haplogroup F are found in some 90% of the world's population, but almost exclusively outside of sub-Saharan Africa.

F xG,H,I,J,K is rare in modern populations and peaks in South Asia, especially Sri Lanka.[10] It also appears to have long been present in South East Asia; it has been reported at rates of 4–5% in Sulawesi and Lembata. One study, which did not comprehensively screen for other subclades of F-M89 (including some subclades of GHIJK), found that Indonesian men with the SNP P14/PF2704 (which is equivalent to M89), comprise 1.8% of men in West Timor, 1.5% of Flores 5.4% of Lembata 2.3% of Sulawesi and 0.2% in Sumatra.[15][16] F* (F xF1,F2,F3) has been reported among 10% of males in Sri Lanka and South India, 5% in Pakistan, as well as lower levels among the Tamang people (Nepal), and in Iran. F1 (P91), F2 (M427) and F3 (M481; previously F5) are all highly rare and virtually exclusive to regions/ethnic minorities in Sri Lanka, India, Nepal, South China, Thailand, Burma, and Vietnam. In such cases, however, the possibility of misidentification is considered to be relatively high and some may belong to misidentified subclades of Haplogroup GHIJK.[17]

Haplogroup G (M201)

Haplogroup G (M201) originated some 48,000 years ago and its most recent common ancestor likely lived 26,000 years ago in the Middle East. It spread to Europe with the Neolithic Revolution.

It is found in many ethnic groups in Eurasia; most common in the Caucasus, Iran, Anatolia and the Levant. Found in almost all European countries, but most common in Gagauzia, southeastern Romania, Greece, Italy, Spain, Portugal, Tyrol, and Bohemia with highest concentrations on some Mediterranean islands; uncommon in Northern Europe.[18][19]

G-M201 is also found in small numbers in northwestern China and India, Bangladesh, Pakistan, Sri Lanka, Malaysia, and North Africa.

  • Haplogroup G1
  • Haplogroup G2
    • Haplogroup G2a
      • Haplogroup G2a1
      • Haplogroup G2a2
      • Haplogroup G2a3
        • Haplogroup G2a3a
        • Haplogroup G2a3b
          • Haplogroup G2a3b1
      • Haplogroup G2b
      • Haplogroup G2c (formerly Haplogroup G5)
        • Haplogroup G2c1

Haplogroup H (M69)
Main article: Haplogroup H (Y-DNA)

Haplogroup H (M69) probably emerged in South Central Asia or South Asia, about 48,000 years BP, and remains largely prevalent there in the forms of H1 (M69) and H3 (Z5857). Its sub-clades are also found in lower frequencies in Iran, Central Asia, across the middle-east, and the Arabian peninsula.

However, H2 (P96) is present in Europe since the Neolithic and H1a1 (M82) spread westward in the Medieval era with the migration of the Roma people.

Haplogroup I (M170)

Haplogroup I (M170, M258) is found mainly in Europe and the Caucasus.

  • Haplogroup I1 Nordid/Nordic Euopids (M253) Found mainly in northern Europe
  • Haplogroup I2 Dinarid/Dinaric Europids (P215) Found mainly in Balkans, southeast Europe and Sardinia save for I2B1 (m223) which is found at a moderate frequency in Western, Central, and Northern Europe.

Haplogroup J (M304)
Main article: Haplogroup J (Y-DNA)

Haplogroup J (M304, S6, S34, S35) is found mainly in the Middle East and South-East Europe.

Haplogroup K (M9)

Haplogroup K (M9) is spread all over Eurasia, Oceania and among Native Americans.

K(xLT,K2a,K2b) – that is, K*, K2c, K2d or K2e – is found mainly in Melanesia, Aboriginal Australians, India, Polynesia and Island South East Asia.

Haplogroups L and T (K1)
Main article: Haplogroup LT

Haplogroup L (M20) is found in South Asia, Central Asia, South-West Asia, and the Mediterranean.

Haplogroup T (M184, M70, M193, M272) is found at high levels in the Horn of Africa (mainly Cushitic-speaking peoples), parts of South Asia, the Middle East, and the Mediterranean. T-M184 is also found in significant minorities of Sciaccensi, Stilfser, Egyptians, Omanis, Sephardi Jews,[20] Ibizans (Eivissencs), and Toubou. It is also found at low frequencies in other parts of the Mediterranean and South Asia.

Haplogroup K2 (K-M526)
Main article: Haplogroup K2

The only living males reported to carry the basal paragroup K2* are indigenous Australians. Major studies published in 2014 and 2015 suggest that up to 27% of Aboriginal Australian males carry K2*, while others carry a subclade of K2.

Haplogroups K2a, K2a1, NO & NO1
Main article: Haplogroup K2a (Y-DNA)

Haplogroup N

Haplogroup N (M231) is found through northern Eurasia, especially among speakers of the Uralic languages.

Haplogroup N possibly originated in eastern Asia and spread both northward and westward into Siberia, being the most common group found in some Uralic-speaking peoples.

Haplogroup O

Haplogroup O (M175) is found with its highest frequency in East Asia and Southeast Asia, with lower frequencies in the South Pacific, Central Asia, South Asia, and islands in the Indian Ocean (e.g. Madagascar, the Comoros).

Haplogroups K2b1, M & S
Main article: Haplogroup K2b1 (Y-DNA)

No examples of the basal paragroup K2b1* have been identified. Males carrying subclades of K2b1 are found primarily among Papuan peoples, Micronesian peoples, indigenous Australians, and Polynesians.

Its primary subclades are two major haplogroups:

  • Haplogroup S (B254) also known as K2b1a: found in the highlands of Papua New Guinea and;
  • Haplogroup M (P256) also known as K2b1b: found in New Guinea and Melanesia.

Haplogroup P (K2b2)
Main article: Haplogroup P (Y-DNA)

Haplogroup P (P295) has two primary branches: P1 (P-M45) and the extremely rare P2 (P-B253).[21]

P*, P1* and P2 are found together only on the island of Luzon, in The Philippines.[21] In particular, P* and P1* are found at significant rates among members of the Aeta (or Agta) people of Luzon.[22] While, P1* is now more common among living individuals in Eastern Siberia and Central Asia, it is also found at low levels in mainland South East Asia and South Asia. Considered together, these distributions tend to suggest that P* emerged from K2b in South East Asia.[22][23]

P1 is also the parent node of two primary clades:

  • Haplogroup Q (Q-M242) and;
  • Haplogroup R (R-M207). These share the common marker M45 in addition to at least 18 other SNPs.

Haplogroup Q (MEH2, M242, P36) found in Siberia and the Americas Haplogroup R (M207, M306): found in Europe, West Asia, Central Asia, and South Asia

Haplogroup Q M242

Q is defined by the SNP M242. It is believed to have arisen in Central Asia approximately 32,000 years ago.[24][25] The subclades of Haplogroup Q with their defining mutation(s), according to the 2008 ISOGG tree[26] are provided below. ss4 bp, rs41352448, is not represented in the ISOGG 2008 tree because it is a value for an STR. This low frequency value has been found as a novel Q lineage (Q5) in Indian populations[27]

The 2008 ISOGG tree

Haplogroup R (M207)
Main article: Haplogroup R (Y-DNA)
The hypothetical divergence of Haplogroup R and its descendants.

Haplogroup R is defined by the SNP M207. The bulk of Haplogroup R is represented in descendant subclade R1 (M173), which likely originated on the Eurasian Steppes. R1 has two descendant subclades: R1a and R1b.

R1a is associated with the proto-Indo-Iranian and Balto-Slavic peoples, and is now found primarily in Central Asia, South Asia, and Eastern Europe.

Haplogroup R1b is the dominant haplogroup of Western Europe and also found sparsely distributed among various peoples of Asia and Africa. Its subclade R1b1a2 (M269) is the haplogroup that is most commonly found among modern Western European populations, and has been associated with the Italo-Celtic and Germanic peoples.

  • Haplogroup R1 (M173) Found throughout western Eurasia
    • Haplogroup R1a (M420) Found in Central Asia, South Asia, and Central, Northern and Eastern Europe, Balkans
    • Haplogroup R1b (M343) Found in Western Europe, West Asia, Central Asia, North Africa, and northern Cameroon
  • Haplogroup R2 (M124) Found in South Asia, Caucasus, Central Asia, and Eastern Europe

Chronological development of haplogroups
Haplogroup Possible time of origin Possible place of origin Possible TMRCA[33][12]
A00235,900[4] or 275,000 years ago[34]Africa[35]235,900 years ago
BT130,700 years ago[4]Africa88,000 years ago
CT88,000[4] or 101-100,000 years ago[36][37]Africa68,500 years ago
E65,200,[4] 69,000,[38] or 73,000 years ago[39]East Africa[40] or Asia[15]53,100 years ago
F65,900 years ago[4]Eurasia48,800 years ago
G48,500 years ago[4]Middle East26,200 years ago
IJ47,200 years ago[4]Middle East42,900 years ago
K47,200 years ago[4]Asia45,400 years ago
P45,400 years ago[4]Asia31,900 years ago
J42,900 years ago[4][41]Middle East31,600 years ago
I42,900 years ago[4]Europe27,500 years ago
E-M215 (E1b1b)42,300 years ago[4][42]East Africa34,800 years ago
E-V38 (E1b1a)42,300 years ago[4][42]East Africa40,100 years ago
N36,800 years ago[4][43]Asia22,100 years ago
E1b1b-M3534,800 years ago[4][42]East Africa24,100 years ago
R31,900 years ago[4]Asia28,200 years ago
J-M267 (J1)31,600 years ago[4][41]Middle East18,500 years ago
J-M172 (J2)31,600 years ago[4][41]Middle East27,800 years ago[4][44]
R-M173 (R1)28,200 years ago[4]Asia22,800 years ago
I-M253 (I1)27,500 years ago[4][45][46]Europe4,600 years ago
I-M438 (I2)27,500 years ago[4][46]Europe21,800 years ago
R-M420 (R1a)22,800 years ago[4][47]Eurasia18,300 years ago
R-M343 (R1b)22,800 years ago[4][48]Eurasia[49]20,400 years ago
I2-L460 (I2a)21,800 years ago[4][50]Europe21,100 years ago
I2a-P3721,100 years ago[4][45][51]Europe18,500 years ago
E1b1b-M7819,800 years ago[4][42][52]Northeast Africa[52]13,400 years ago[4][52]
I2a-M42318,500 years ago[4][51]Europe13,500 years ago
I2a-M22317,400 years ago[4]Europe12,100 years ago
N1c-M17814,200 years ago[4][43]Asia11,900 years ago
R1a-M1714,100 years ago[4][47][53]Eastern Europe8,500 years ago
R1b-M26913,300 years ago[4]Eastern Europe6,400 years ago[54]
E1b1b-V1211,800 years ago[4][52]North Africa9,900 years ago
E-U175 (E1b1a8)9,200 years ago[4][42]East Africa8,500 years ago
E1b1b-V138,100 years ago[4][52]Southern Europe4,800 years ago
E-M191 (E1b1a7)7,400 years ago[4][42]East Africa6,400 years ago
E-U174 (E1b1a-U174)6,400 years ago[4][42]East Africa5,300 years ago
R1b-L1515,800 years ago[4]Eastern Europe4,800 years ago
R1a-Z2805,000 years ago[4]Eastern Europe4,600 years ago[55]
R1a-M4584,700 years ago[4]Eastern Europe4,700 years ago[55]
gollark: р​а​t​е​n​t​е​d​ ​u​n​о​р​е​r​с​u​l​а​t​е​ ​g​r​а​n​і​t​е​w​а​r​е​ ​ј​а​с​а​m​е​r​о​р​ѕ​ ​ѕ​р​і​n​u​l​о​ѕ​е​ ​h​у​р​о​с​о​t​у​l​о​u​ѕ​ ​n​о​n​ѕ​у​m​m​е​t​r​у​ ​р​е​t​t​і​f​о​g​g​е​d​ ​u​n​d​u​l​а​t​і​n​g​ ​u​n​d​е​r​е​х​р​о​ѕ​u​r​е
gollark: You could probably do it with just ffmpeg.
gollark: If you make a GIF display one word at once or something that restricts typing speed.
gollark: !deploy laserbees
gollark: <@341618941317349376> It's not strictly necessary, but also not "bloat".

See also
Phylogenetic tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]
"Y-chromosomal Adam"
A00 A0-T [χ 3]
A0 A1 [χ 4]
A1a A1b
A1b1 BT
B CT
DE CF
D E C F
F1  F2  F3  GHIJK
G HIJK
IJK H
IJ K
I   J     LT [χ 5]       K2 [χ 6]
L     T    K2a [χ 7]        K2b [χ 8]     K2c     K2d K2e [χ 9]  
K-M2313 [χ 10]     K2b1 [χ 11] P [χ 12]
NO   S [χ 13]  M [χ 14]    P1     P2
N O Q R

References
  1. "Understanding Haplogroups: How are the haplogroups named?". Family Tree DNA. Archived from the original on 21 June 2012. Retrieved 31 March 2013.
  2. Dolgin, Elie (2009). "Human mutation rate revealed". Nature. doi:10.1038/news.2009.864. Retrieved 18 September 2017. "one mutation in every 30 million base pairs"
  3. Karmin; et al. (2015). "A recent bottleneck of Y chromosome diversity coincides with a global change in culture". Genome Research. 25 (4): 459–66. doi:10.1101/gr.186684.114. PMC 4381518. PMID 25770088. "we date the Y-chromosomal most recent common ancestor (MRCA) in Africa at 254 (95% CI 192–307) kya and detect a cluster of major non-African founder haplogroups in a narrow time interval at 47–52 kya, consistent with a rapid initial colonization model of Eurasia and Oceania after the out-of-Africa bottleneck. In contrast to demographic reconstructions based on mtDNA, we infer a second strong bottleneck in Y-chromosome lineages dating to the last 10 ky. We hypothesize that this bottleneck is caused by cultural changes affecting variance of reproductive success among males."
  4. "YFull YTree". YFull. Retrieved 15 September 2017.
  5. {{cite web|title=Something Weird Happened to Men 7,000 Years Ago, And We Finally Know Why | url=https://www.sciencealert.com/neolithic-y-chromosome-bottleneck-warring-patrilineal-clans | quote=Around 7,000 years ago - all the way back in the Neolithic - something really peculiar happened to human genetic diversity. Over the next 2,000 years, and seen across Africa, Europe and Asia, the genetic diversity of the Y chromosome collapsed, becoming as though there was only one man for every 17 women.
  6. "Understanding Results: Y-DNA Single Nucleotide Polymorphism (SNP): What is a Y-chromosome DNA (Y-DNA) haplogroup?". Family Tree DNA. Retrieved 31 March 2013. Y-chromosome DNA (Y-DNA) haplogroups are the major branches on the human paternal family tree. Each haplogroup has many subbranches. These are subclades.
  7. "myFTDNA 2.0 User Guide: Y-DNA: What is the Y-DNA – Matches page?". Family Tree DNA. Retrieved 31 March 2013. A terminal SNP determines the terminal (final) subbranch on the Y-DNA Tree to which someone belongs.
  8. "Understanding Results: Y-DNA Single Nucleotide Polymorphism (SNP): How are haplogroups and their subclades named?". Family Tree DNA. Retrieved 31 March 2013.
  9. "Family Tree DNA – Genetic Testing for Ancestry, Family History & Genealogy". familytreedna.com.
  10. Copyright 2015 ISOGG. "ISOGG 2015 Y-DNA Haplogroup Tree Trunk". isogg.org.
  11. Underhill and Kivisild; Kivisild, T (2007). "Use of Y Chromosome and Mitochondrial DNA Population Structure in Tracing Human Migrations". Annu. Rev. Genet. 41 (1): 539–64. doi:10.1146/annurev.genet.41.110306.130407. PMID 18076332.
  12. Karafet, TM; Mendez, FL; Meilerman, MB; Underhill, PA; Zegura, SL; Hammer, MF (2008). "New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree". Genome Research. 18 (5): 830–38. doi:10.1101/gr.7172008. PMC 2336805. PMID 18385274.
  13. Kamin M, Saag L, Vincente M, et al. (April 2015). "A recent bottleneck of Y chromosome diversity coincides with a global change in culture". Genome Research. 25 (4): 459–66. doi:10.1101/gr.186684.114. PMC 4381518. PMID 25770088.
  14. Haber M, Jones AL, Connel BA, Asan, Arciero E, Huanming Y, Thomas MG, Xue Y, Tyler-Smith C (June 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): 1421–28. doi:10.1534/genetics.119.302368. PMC 6707464. PMID 31196864.
  15. Chiaroni, Jacques; Underhill, Peter A.; Cavalli-Sforza, Luca L. (1 December 2009). "Y chromosome diversity, human expansion, drift, and cultural evolution". Proceedings of the National Academy of Sciences of the United States of America. 106 (48): 20174–79. Bibcode:2009PNAS..10620174C. doi:10.1073/pnas.0910803106. PMC 2787129. PMID 19920170.
  16. Tumonggor, Meryanne K (2014). "Isolation, contact and social behavior shaped genetic diversity in West Timor". Journal of Human Genetics. 59 (9): 494–503. doi:10.1038/jhg.2014.62. PMC 4521296. PMID 25078354.
  17. This was, for instance, the case with the original subclade F3 (M96), which has since been renamed Haplogroup H2.
  18. Passarino G, Cavalleri GL, Lin AA, Cavalli-Sforza LL, Børresen-Dale AL, Underhill PA (2002). "Different genetic components in the Norwegian population revealed by the analysis of mtDNA and Y chromosome polymorphisms". European Journal of Human Genetics. 10 (9): 521–29. doi:10.1038/sj.ejhg.5200834. PMID 12173029.
  19. Karlsson, Andreas O; Wallerström, Thomas; Götherström, Anders; Holmlund, Gunilla (2006). "Y-chromosome diversity in Sweden – A long-time perspective". European Journal of Human Genetics. 14 (8): 963–70. doi:10.1038/sj.ejhg.5201651. PMID 16724001.
  20. Nogueiro, Inês (2009). "Phylogeographic analysis of paternal lineages in NE Portuguese Jewish communities". American Journal of Physical Anthropology. 141 (3): 373–81. doi:10.1002/ajpa.21154. PMID 19918998.
  21. ISOGG, 2016, Y-DNA Haplogroup P and its Subclades – 2016 (20 June 2016).
  22. Tumonggor, Meryanne K; Karafet, Tatiana M; Downey, Sean; Lansing, J Stephen; Norquest, Peter; Sudoyo, Herawati; Hammer, Michael F; Cox, Murray P (31 July 2014). "Isolation, contact and social behavior shaped genetic diversity in West Timor". Journal of Human Genetics. 59 (9): 494–503. doi:10.1038/jhg.2014.62. PMC 4521296. PMID 25078354.
  23. Tatiana M Karafet; et al. (2015). "Improved phylogenetic resolution and rapid diversification of Y-chromosome haplogroup K-M526 in Southeast Asia". European Journal of Human Genetics. 23 (3): 369–73. doi:10.1038/ejhg.2014.106. PMC 4326703. PMID 24896152.
  24. Fagundes, Nelson J.R.; Ricardo Kanitz; Roberta Eckert; Ana C.S. Valls; Mauricio R. Bogo; Francisco M. Salzano; David Glenn Smith; Wilson A. Silva; Marco A. Zago; Andrea K. Ribeiro-dos-Santos; Sidney E.B. Santos; Maria Luiza Petzl-Erler; Sandro L. Bonatto (2008). "Mitochondrial Population Genomics Supports a Single Pre-Clovis Origin with a Coastal Route for the Peopling of the Americas" (PDF). American Journal of Human Genetics. 82 (3): 583–92. doi:10.1016/j.ajhg.2007.11.013. PMC 2427228. PMID 18313026. Archived from the original (PDF) on 2009-03-25. Retrieved 2013-05-22. Since the first studies, it has been found that extant Native American populations exhibit almost exclusively five "mtDNA haplogroups" (A–D and X)6 classified in the autochthonous haplogroups A2, B2, C1, D1, and X2a.7 Haplogroups A–D are found all over the New World and are frequent in Asia, supporting a northeastern Asian origin of these lineages
  25. Zegura, S. L.; Karafet, TM; Zhivotovsky, LA; Hammer, MF (2003). "High-Resolution SNPs and Microsatellite Haplotypes Point to a Single, Recent Entry of Native American Y Chromosomes into the Americas". Molecular Biology and Evolution. 21 (1): 164–75. doi:10.1093/molbev/msh009. PMID 14595095.
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