Haplogroup K2

Haplogroup K2
Possible time of origin	47,000-50,000 years BP[1]
Possible place of origin	Southeast[2] or East Asia[3]
Ancestor	K
Descendants	K2a (M2308); K2b (MPS); K2c; K2d; K2e.
Defining mutations	rs2033003 (M526)

Haplogroup K2, also known as K-M526 and formerly known as K(xLT) and MNOPS,[4] is a human Y-DNA haplogroup.

Relative to its age, the internal structure of K2 is extremely complex, and subclades of it are carried by males native to regions including Australasia, Oceania, Southeast Asia, South Asia, East Asia, Central Asia, the Americas, Europe, and the Horn of Africa. Many of its branches are very common, the most numerically important being R in Europe and South Asia and O in East and Southeast Asia (as well as recent immigrants to other continents). Haplogroups N and Q, while they are less common overall, are also very widespread, in northern Eurasia and the Americas respectively. M and S are almost entirely restricted to Oceania and eastern Indonesia, where they occur at high frequency.

Rare subclades outside of these major lineages are known mainly from Island Southeast Asia (including the Andaman Islands and the Philippines).

Basal paragroup K2* has not been identified among living males or ancient remains.

K2a* has been found only in Upper Paleolithic remains from western Siberia and the Balkans, known respectively as "Ust'-Ishim man and "Oase-1'.".[5] The only primary branch of K2a, known as K-M2313*, has been documented in two living individuals, who have ethnic ties to South Asia and South East Asia respectively: a Telugu from India and an ethnic Malay from Singapore. In addition, K-Y28299, which appears to be a primary branch of K-M2313, has been found in three living individuals from India.[6] Another subclade, NO (M214)* – which for a time was thought to be synonymous with K2a (M2308)* – has not been identified in living individuals or remains.

Basal paragroup K2b* has not been identified among living males but was found in Upper Paleolithic Tianyuan man from China.[7] K2b1 (P397/P399) known previously as Haplogroup MS, and Haplogroup P (P-P295), also known as K2b2 are the only primary clades of K2b. The population geneticist Tatiana Karafet and other researchers (2014) point out that both K2b1 and P* are virtually restricted geographically to South East Asia and Oceania.[2] Whereas, in a striking contrast, P1 (P-M45) and its primary subclades Q and R now make up "the most frequent haplogroup in Europe, the Americas, and Central Asia and South Asia". According to Karafet et al., the estimated dates for the branching of K, K2, K2b and P point to a "rapid diversification" within K2 "that likely occurred in Southeast Asia", with subsequent "westward expansions" of P*, P1, Q and R.[2]

Structure

A direct descendant of Haplogroup K, K2 is a sibling of basal/paragroup K* and Haplogroup LT (also known as K1).

As of 2017, the phylogeny of haplogroup K2 is as follows:

K-M526 (K2) M526 – formerly known as K(xLT) and MNOPS

K-M2308 (K2a) M2308 – found only in ancient remains; see above)[5]
- K-M2313 (K2a1) M2313[5]
  - NO-F549 (NO*) or K-M2335 F549/S22380/M2335/V4208; M2335/Z4952/M2339/E482; CTS11667[6][5]
    - NO1-M214 (NO1*) or NO-F176 M214/Page39; F176/M2314; CTS5858/M2325/F346; CTS11572[6][5][8]
      - N-M231 (N*) M231; CTS2947/M2175; Z4891; CTS10118[6]
      - O-M175 (O*) M175/P186/P191/P196; F369/M1755; F380/M1757/S27659[6]
  - K-Y28299 (primary subclade of K2a; no phylogenetic name as of 2017) Y28299/Y28355; Y28357; Y28412 – found in one living individual in India[6]
  - K-Y28301 (primary subclade of K2a; no phylogenetic name as of 2017) Y28301/Y28328; Y28358; Y28410 – found in two living individuals in India[6]
K-P331 (K2b) M1221/P331/PF5911, CTS2019/M1205, PF5990/L405, PF5969 – subclades of K2b include the major haplogroups M; S, P, Q, and R
K-P261 (K2c) P261
K-P261 (K2d) P402
K-P261 (K2e) M147

Distribution

At the level of highly-derived subclades, K2 is almost universal in some modern Eurasian and Native American populations. Haplogroup NO* (descendant of K2a) includes most males among Southeast Asian, East Asian, and Finno-Ugric populations.[5] The descendants of K2b include the major haplogroups M; S, P, Q, and R. These are now numerically in dominant in: Oceania, Central Asia, Siberia, among Native American populations, Europe, and South Asia.

A rapid diversification within and from K2 (M526), most likely in Southeast Asia, is suggested by estimates of the point in time that K2 branched off from K* (M9). Likewise the branching from K2 of K2b (P331) and Haplogroup P (K2b2 P295) from K2b, as well as Haplogroups Q and R from P (K2b2), and their subsequent expansions westward in Europe,[2] and eastward into the Americas.

K2c, K2d, and K2e are extremely rare subhaplogroups that are found in specific parts of South and Southeast Asia.[9] K2c (P261) has been reported only among males in Bali and K2d (P402) only in Java. K2e (M147), which has been found in two modern cases from South India,[9] was provisionally named "pre-NO" (among other names), as it was believed initially to be ancestral to K2a (NO). However, it was later found to be a primary branch of Haplogroup K2 (K-M526) and a sibling of K2a; the new clade was renamed K2e.

Studies published in 2014 and 2015 found that up to 27% of Aboriginal Australian males carry K-M526*, which could not be classified into a known subclade at the time, and another 27% probably have K2b1a1 (P60, P304, P308; also known as "S-P308") and perhaps 2.0% have Haplogroup M1 – also known as M-M4 (or "M-M186") and K2b1d1.[10][2]

Naming

The name K2 was introduced in 2014, following dissatisfaction with the previous names.

K(xLT), the name introduced by the Y Chromosome Consortium in 2012 to replace MNOPS, was controversial. Under the previous methodology, a term such as "K(xLT)" designated all clades and subclades that belonged to K, but did not belong to Haplogroup LT; the haplogroups subordinate to MNOPS would likely have been renamed "U", "V", "W" and "X", and MNOPS would therefore have become "MNOPSUVWX". This posed a problem, because there was no way to disambiguate between "K(xLT)" in the broad and narrow meanings of the term.

Footnotes

Karafet et al. 2014
https://link.springer.com/article/10.1007/s00439-020-02204-9
Chiaroni, Jacques; Underhill, Peter A.; Cavalli-Sforza, Luca L. (2009). "Y chromosome diversity, human expansion, drift, and cultural evolution". PNAS. 106 (48): 20174–9. Bibcode:2009PNAS..10620174C. doi:10.1073/pnas.0910803106. PMC 2787129. PMID 19920170.
G. David Poznik, Yali Xue, Fernando L. Mendez, et al., 2016, "Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences." Nature Genetics vol. 48, no. 6 (June): pp. 593–599. doi:10.1038/ng.3559.
YFull YTree v5.08, 2017, "K-M2335" (9 December 2017); PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (9 December 2017); GeneticHomeland.com, 2016, DNA Marker Index Chromosome Y V4208 (9 December 2017).
"Downloadable genotypes of present-day and ancient DNA data (compiled from published papers) | David Reich Lab". reich.hms.harvard.edu. Retrieved 2019-09-11.
ISOGG 2016 Y-DNA Haplogroup Tree 2016 (2 August 2016).
Tatiana M. Karafet, Fernando L. Mendez, Herawati Sudoyo, J. Stephen Lansing and Michael F. Hammer; 2015, "Improved phylogenetic resolution and rapid diversification of Y-chromosome haplogroup K-M526 in Southeast Asia", European Journal of Human Genetics, no. 23 (March), pp. 369–73.
Nagle, N. et al., 2015, "Antiquity and diversity of aboriginal Australian Y-chromosomes", American Journal of Physical Anthropology (epub ahead of print version; abstract). (The authors assumed 56% of the samples taken to be non-indigenous.)

Phylogenetic tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]

A1b

A1b1

F1

F2

F3

GHIJK

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

Footnotes

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.
International Society of Genetic Genealogy (ISOGG; 2015), Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.)
Haplogroup A0-T is also known as A-L1085 (and previously as A0'1'2'3'4).
Haplogroup A1 is also known as A1'2'3'4.
Haplogroup LT (L298/P326) is also known as Haplogroup K1.
Between 2002 and 2008, Haplogroup T-M184 was known as "Haplogroup K2". That name has since been re-assigned to K-M526, the sibling of Haplogroup LT.
Haplogroup K2a (M2308) and its primary subclade K-M2313 were separated from Haplogroup NO (F549) in 2016. (This followed the publication of: Poznik GD, Xue Y, Mendez FL, et al. (2016). "Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences". Nature Genetics. 48 (6): 593–9. doi:10.1038/ng.3559. PMC 4884158. PMID 27111036. In the past, other haplogroups, including NO (M214) and K2e had also been identified with the name "K2a".
Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.
Haplogroup K2e (K-M147) was previously known as "Haplogroup X" and "K2a" (but is a sibling subclade of the present K2a).
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: Poznik op. 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)
Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure.
Haplogroup P (P295) is also klnown as K2b2.
Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)
Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)

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[karafet_2014-2] Karafet et al. 2014

[3] ttps://link.springer.com/article/10.1007/s00439-020-02204-9

[Chiaroni2009-4] Chiaroni, Jacques; Underhill, Peter A.; Cavalli-Sforza, Luca L. (2009). "Y chromosome diversity, human expansion, drift, and cultural evolution". PNAS. 106 (48): 20174–9. Bibcode:2009PNAS..10620174C. doi:10.1073/pnas.0910803106. PMC 2787129. PMID 19920170.

[poznik-5] G. David Poznik, Yali Xue, Fernando L. Mendez, et al., 2016, "Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences." Nature Genetics vol. 48, no. 6 (June): pp. 593–599. doi:10.1038/ng.3559.

[F549-6] YFull YTree v5.08, 2017, "K-M2335" (9 December 2017); PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (9 December 2017); GeneticHomeland.com, 2016, DNA Marker Index Chromosome Y V4208 (9 December 2017).

[7] "Downloadable genotypes of present-day and ancient DNA data (compiled from published papers) | David Reich Lab". reich.hms.harvard.edu. Retrieved 2019-09-11.

[ISOGG2016-8] ISOGG 2016 Y-DNA Haplogroup Tree 2016 (2 August 2016).

[Karafet2015-9] Tatiana M. Karafet, Fernando L. Mendez, Herawati Sudoyo, J. Stephen Lansing and Michael F. Hammer; 2015, "Improved phylogenetic resolution and rapid diversification of Y-chromosome haplogroup K-M526 in Southeast Asia", European Journal of Human Genetics, no. 23 (March), pp. 369–73.

[nagle-10] Nagle, N. et al., 2015, "Antiquity and diversity of aboriginal Australian Y-chromosomes", American Journal of Physical Anthropology (epub ahead of print version; abstract). (The authors assumed 56% of the samples taken to be non-indigenous.)

[vanOven2014-11] 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.

[ISOGG2015-12] International Society of Genetic Genealogy (ISOGG; 2015), Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.)

[A0-T-13] Haplogroup A0-T is also known as A-L1085 (and previously as A0'1'2'3'4).

[A1-14] Haplogroup A1 is also known as A1'2'3'4.

[LT-15] Haplogroup LT (L298/P326) is also known as Haplogroup K1.

[K2-16] Between 2002 and 2008, Haplogroup T-M184 was known as "Haplogroup K2". That name has since been re-assigned to K-M526, the sibling of Haplogroup LT.

[K2a-17] Haplogroup K2a (M2308) and its primary subclade K-M2313 were separated from Haplogroup NO (F549) in 2016. (This followed the publication of: Poznik GD, Xue Y, Mendez FL, et al. (2016). "Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences". Nature Genetics. 48 (6): 593–9. doi:10.1038/ng.3559. PMC 4884158. PMID 27111036. In the past, other haplogroups, including NO (M214) and K2e had also been identified with the name "K2a".

[K2b-18] Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.

[K2e-19] Haplogroup K2e (K-M147) was previously known as "Haplogroup X" and "K2a" (but is a sibling subclade of the present K2a).

[K-M2313-20] 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: Poznik op. 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)

[K2b1-21] Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure.

[P-22] Haplogroup P (P295) is also klnown as K2b2.

[S-23] Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)

[M-24] Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)