Haplogroup R1b

Haplogroup R1b
Possible place of origin Western Asia[1]
Ancestor R1
Descendants

One immediate descendant:

  • R1b1 (L278, M415, P25). Two secondary descendants:
    • R1b1a (L754, PF6269, YSC0000022)
    • R1b1b (PH155)
Defining mutations M343

Haplogroup R1b (R-M343), also known as Hg1 and Eu18, is a human Y-chromosome haplogroup.

It is the most frequently occurring paternal lineage in Western Europe, as well as some parts of Russia (e.g. the Bashkir minority) and Central Africa (e.g. Chad and Cameroon). The clade is also present at lower frequencies throughout Eastern Europe, Western Asia, as well as parts of North Africa and Central Asia.

R1b also reaches high frequencies in the Americas and Australasia, due largely to immigration from Western Europe. There is an ongoing debate regarding the origins of R1b subclades found at significant levels among some indigenous peoples of the Americas, such as speakers of Algic languages in central Canada.

R1b has one primary branch, R1b1 (L278), which in turn has two primary branches: R1b1a (L754) and R1b1b (PH155). R1b1a is found mostly in Western Europe, although the Fula and Chadic-speaking peoples of Africa are dominated by R1b1a2 (PF6279/V88).

R1b1b (PH155) is so rare and widely dispersed that it is difficult to draw any conclusions about its origins. It has been found in Bahrain, Bhutan, Tajikistan, Turkey and Western China.

Western Europe is dominated by the downstream subclades of R1b1a – especially R1b1a1a2 (R-M269; known previously as R1b1a2).

The oldest human remains found to carry with R1b so far are:

Origin and dispersal

R1b is a subclade within the "macro-haplogroup" Haplogroup K (K-M9), which is one of the predominant groupings of all the rest of human male lines outside of Africa. K* is believed to have originated in Asia (as is the case with an even earlier ancestral haplogroup, F (F-M89). Karafet T. 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." [2]

Three autosomal genetic studies in 2015 gave support to the Kurgan hypothesis of Gimbutas regarding the proto-Indo-European homeland. 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 Russian 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.[3][4][5]

The point of origin of R1b is thought to lie in Eurasia, most likely in Western Asia.[6] T. Karafet et al. (2008) estimated the age of R1, the parent of R1b, as 18,500 years before present.[7]

Early research into the origins of R1b focused on Europe. In 2000, Ornella Semino and colleagues argued that R1b had been in Europe before the end of the Ice Age, and had spread north from an Iberian refuge after the Last Glacial Maximum.[8] Age estimates of R1b in Europe have steadily decreased in more recent studies, at least concerning the majority of R1b, with more recent studies suggesting a Neolithic age or younger.[6][9][10][11] On the other hand, Morelli et al. have recently (in 2010) attempted to defend a Palaeolithic origin for R1b1b2.[12] Irrespective of microsatellite coalescence calculations, Chikhi et al. pointed out that the timing of molecular divergences does not coincide with population splits; the TMRCA of haplogroup R1b (whether in the Palaeolithic or Neolithic) dates to its point of origin somewhere in Eurasia, and not its arrival in western Europe.[13] Summing up, Michael R. Maglio argues that the closest branch of R1b is from Iberia and its small subclades found in West Asia, the Near East and Africa are examples of back migration, and not of its origin.[14]

However, as Barbara Arredi and colleagues were the first to point out, the distribution of R1b microsatellite variance in Europe forms a cline from east to west, which is more consistent with an entry into Europe from Western Asia with the spread of farming.[11] A 2009 paper by Chiaroni et al. added to this perspective by using R1b as an example of a wave haplogroup distribution, in this case from east to west.[15] The proposal of a southeastern origin of R1b were supported by three detailed studies based on large datasets published in 2010. These detected that the earliest subclades of R1b are found in western Asia and the most recent in western Europe.[6][9][16]

While age estimates in these articles are all more recent than the Last Glacial Maximum, all mention the Neolithic, when farming was introduced to Europe from the Middle East as a possible candidate period. Myres et al. (August 2010), and Cruciani et al. (August 2010) both remained undecided on the exact dating of the migration or migrations responsible for this distribution, not ruling out migrations as early as the Mesolithic or as late as the Hallstatt culture but more probably Late Neolithic.[6] They noted that direct evidence from ancient DNA may be needed to resolve these gene flows.[6] Lee et al. (May 2012) analysed the ancient DNA of human remains from the Late Neolithic Beaker culture site of Kromsdorf, Germany identifying two males as belonging to the Y haplogroup R1b.[17] Analysis of ancient Y DNA from the remains of populations derived from early Neolithic Central and North European Linear Pottery culture settlements have not yet found males belonging to haplogroup R1b.[18][19]

The suggestion has also been made that the spread of R1b in Western Europe may coincide with the spread of the Centum branch of the Indo-European languages during the early Bronze Age.[20]

One of the highest levels of R1b is found among the Basques, who speak a non- Indo-European language isolate; which contradicts an exclusively or predominantly Indo-European source. One hypothesis about the case of the Basques is that a male-dominated Indo-European-speaking people invaded and conquered the Basque region, and then, having brought no or few women with them, married local women, possibly from a matrilineal society. The women then raised the children that resulted to speak their own language and with their cultural practices, rather than those of the fathers'. This possible explanation further addresses the point that while other high-R1b regions in Western Europe (such as the British Isles and southern Germany) show disproportionately high incidences of MtDNA haplogroups that correspond to a Pontic Steppes origin (specifically MtDNA Haplogroups I, U2, U3, U4, and W), the Basque region does not. In fact, the Basque region displays virtually no MtDNA for which Pontic Steppes origin could be claimed.[20]

In the 2016 Nature article 'The genetic history of Ice Age Europe', an individual from the Epigravettian cultural context in Italy (Villabruna) is mentioned, who lived circa 12,000 BC and reportedly belonged to Y-DNA group R1b1a (L754).

Structure

External phylogeny

R1b is a part of the broader Haplogroup K-M9 and its linear descendants K2, K2b and P, which is also known as K2b2.

Karafet et al. (2014) suggest that a "rapid diversification process of K-M526 likely occurred in Southeast Asia, with subsequent westward expansions of the ancestors of haplogroups R and Q." [2]

Internal structure

Names such as R1b, R1b1 and so on are phylogenetic (i.e. "family tree") names which make clear their place within the branching of haplogroups, or the phylogenetic tree. An alternative way of naming the same haplogroups and subclades refers to their defining SNP mutations: for example, R-M343 is equivalent to R1b.[21] Phylogenetic names change with new discoveries and SNP-based names are consequently reclassified within the phylogenetic tree. In some cases, an SNP is found to be unreliable as a defining mutation and an SNP-based name is removed completely. For example, before 2005, R1b was synonymous with R-P25, which was later reclassified as R1b1; in 2016, R-P25 was removed completely as a defining SNP, due to a significant rate of back-mutation.[22]

This is the basic outline of R1b according to the ISOGG Tree as it stood on January 30, 2017.[1]

Basic phylogenetic tree for R1b
2017 ISOGG tree
 M343/PF6242 


  R-M343* (R1b*). Now virtually restricted to Iran.


 L278 


  R-L278* (R1b1*). Now found at low levels throughout Western Eurasia.


 L754/PF6269/YSC0000022 


  R-L754* (R1b1a*). Found predominantly in Western Eurasia and Africa.


 L388/PF6468/L389/PF6531 


  R-L388* (R1b1a1*). Either extremely rare or extinct.


 P297/PF6398 


R-P297* (R1b1a1a*; previously R1b1a*). Appears to be extinct.


 M73 

R-M73 (R1b1a1a1) – now common only in Siberia and Central Asia.


 M269/PF6517 

R-M269 (R1b1a1a2; previously R1b1a2) – now extremely common throughout Western Europe, but also found at lower levels in many other parts of Western Eurasia and the Mediterranean.



 V1636

  R-V1636 (R1b1a1b). Found in the Mediterranean region at low frequencies.



 PF6279/V88 

  R-V88 (R1b1a2). Now the most common form of R1b in Sub-Saharan Africa. (Previously known as R1b1c.)



PH155

  R-PH155 (R1b1b). Extremely rare; examples have been found in Bahrain, Bhutan, Tajikistan, Turkey, and Xinjiang.




R1b* (R-M343*)

R1b* – that is, males with M343, but no subsequent distinguishing SNP mutations – is extremely rare. The only population yet recorded with a definite significant proportion of R1b* are the Kurds of southeastern Kazakhstan with 13%.[6][23] However, more recently, a large study of Y-chromosome variation in Iran, revealed R1b* as high as 4.3% among Iranian sub-populations.[24]

While studies in 2005–08 suggested that R1b* may occur at high levels in Jordan, Egypt and Sudan, subsequent research indicates that the samples concerned most likely belong to the subclade R1b1a2 (R-V88), which is now concentrated among African populations, following back migration from Asia.[25][26][27][28][29] It remains a possibility that some or even most of these cases may be R1b* (R-M343*), R1a* (R-M420*), an otherwise undocumented branch of R1, and/or back-mutations of a marker, from a positive to a negative ancestral state,[30] constituting, in other words, undocumented subclades of R1b. Thus demonstrating the importance of testing for SNPs critical in identifying subclades.


A compilation of previous studies regarding the distribution of R1b can be found in Cruciani et al. (2010).[31] It is summarised in the table following. (It should be noted that Cruciani did not include some studies suggesting even higher frequencies of R1b1a1a2 [R-M269] in some parts of Western Europe.)

Continent Region Sample size Total R1b R-P25
(unreliable marker for R1b1*) 		R-V88 (R1b1a2) R-M269 (R1b1a1a2) R-M73 (R1b1a1a1)
AfricaNorthern Africa6915.9%0.0%5.2%0.7%0.0%
AfricaCentral Sahel Region46123.0%0.0%23.0%0.0%0.0%
AfricaWestern Africa 1230.0%0.0%0.0%0.0%0.0%
AfricaEastern Africa 4420.0%0.0%0.0%0.0%0.0%
AfricaSouthern Africa 1050.0%0.0%0.0%0.0%0.0%
EuropeWestern Europeans 46557.8%0.0%0.0%57.8%0.0%
EuropeNorth-west Europeans4355.8%0.0%0.0%55.8%0.0%
EuropeCentral Europeans 7742.9%0.0%0.0%42.9%0.0%
EuropeNorth Eastern Europeans741.4%0.0%0.0%1.4%0.0%
EuropeRussians 606.7%0.0%0.0%6.7%0.0%
EuropeEastern Europeans 14920.8%0.0%0.0%20.8%0.0%
EuropeSouth-east Europeans 51013.1%0.0%0.2%12.9%0.0%
AsiaWest Asians 3285.8%0.0%0.3%5.5%0.0%
AsiaSouth Asians 2884.8%0.0%0.0%1.7%3.1%
AsiaSouth-east Asians100.0%0.0%0.0%0.0%0.0%
AsiaNorth-east Asians300.0%0.0%0.0%0.0%0.0%
AsiaEast Asians 1560.6%0.0%0.0%0.6%0.0%
TOTAL5326

R1b1 (R-L278)

R1b1* or R-L278* is rare among modern males. However, it has been found in two skeletons from prehistoric Europe: a male from the Mesolithic Samara culture (a pre-Yamna people who lived immediately north of the Caspian Sea) buried in about 5650–5555 BCE, and a male from the early Neolithic Cardial culture, buried in about 5178–5066 BCE at the Els Trocs site in the Pyrenees (modern Aragon, Spain).[32]

Some examples described in older articles, for example two found in Turkey,[21] are now thought to be mostly in the more recently discovered sub-clade R1b1a2 (R-V88). Most examples of R1b therefore fall into subclades R1b1a2 (R-V88) or R1b1a (R-P297). Cruciani et al. in the large 2010 study found 3 cases amongst 1173 Italians, 1 out of 328 West Asians and 1 out of 156 East Asians.[31] Varzari found 3 cases in the Ukraine, in a study of 322 people from the Dniester-Carpathian Mountains region, who were P25 positive, but M269 negative.[33] Cases from older studies are mainly from Africa, the Middle East or Mediterranean, and are discussed below as probable cases of R1b1a2 (R-V88).

R1b1a (R-L754)

R1b1a (R-L754) was carried by an individual known as Villabruna 1, who lived c. 15,000 years BP in north-west Italy, and belonged to the Epigravettian culture.

Living individuals positive for L761, an equivalent to L754, have been found at high frequencies among the Toubou population inhabiting Chad (34%).[34]

R1b1a1 (R-L388)

R-L388, also known as R1b1a1 (L388/PF6468, L389/PF6531) appears to be rare or extinct in its basal form. Its subclades are also relatively rare and found in various parts of South West Asia, the Mediterranean basin and continental Europe.

R1b1a1a (R-P297)

The SNP marker P297 was recognised in 2008 as ancestral to the significant subclades M73 and M269, combining them into one cluster.[7] This had been given the phylogenetic name R1b1a1a (and, previously, R1b1a).

A majority of Eurasian R1b falls within this subclade, representing a very large modern population. Although P297 itself has not yet been much tested for, the same population has been relatively well studied in terms of other markers. Therefore, the branching within this clade can be explained in relatively high detail below. The skeleton of a male from a Chalcolithic Yamna burial in the Middle-Volga-Samara area, dated to around 3305–2925 BC, was found to possibly contain R1b1a* being P297 positive but L51 negative.[32]

R1b1a1a1 (R-M73)

R-M73 is reportedly the dominant haplogroup among the Kumandin of the Altai Republic in Russia.[35]

While early research into R-M73 claimed that it was significantly represented among the Hazara of Afghanistan and the Bashkirs of the Ural Mountains, this has apparently been overturned. For example, supporting material from a 2010 study by Behar et al. suggested that Sengupta et al. (2006) had misidentified Hazara individuals, who instead belonged to R2 or Q.[36][37][38] Likewise, most Bashkir males have been found to belong to U-152 (R1b1a1a2a1a2b) and some, mostly from south-eastern Bashkortostan, belonged to Haplogroup Q-M25 (Q1a1b) rather than R1b.

R1b1a1a2 (R-M269)
Projected spatial frequency distribution within Europe of haplogroup R-M269.[9]

R-M269, or R1b1a1a2 (as of 2017) amongst other names,[39] is now the most common Y-DNA lineage in European males. It is carried by an estimated 110 million males in Europe.[9]

R-M269 has received significant scientific and popular interest due to its possible connection to the Indo-European expansion in Europe. Specifically the R-L23 (R-Z2103) subclade has been found to be prevalent in ancient DNA associated with the Yamna culture.[32] R-M269 likely originated in Western Asia and was present in Europe by the Neolithic period.[1][6][11][16] The distribution of subclades within Europe is substantially due to the various migrations of the Bronze and Iron Age. Western European populations are divided between the R-P312/S116 and R-U106/S21 subclades of R-M412 (R-L51). In Western Europe it is present but in generally much lower levels apart from "an instance of 27% in Switzerland's Upper Rhone Valley."[6]

Distribution of R-M269 in Europe increases in frequency from east to west. It peaks at the national level in Wales at a rate of 92%, at 82% in Ireland, 70% in Scotland, 68% in Spain, 60% in France (76% in Normandy), about 60% in Portugal, 53% in Italy,[6] 45% in Eastern England, 50% in Germany, 50% in the Netherlands, 42% in Iceland, and 43% in Denmark. R-M269 reaches levels as high as 95% in parts of Ireland. It has also been found at lower frequencies throughout central Eurasia,[40] but with relatively high frequency among the Bashkirs of the Perm region (84.0%).[41] This marker is present in China and India at frequencies of less than one percent. In Africa and adjoining islands, while R-V88 (R1b1a2) is more strongly represented, R-M269 appears to have been present at very high rates since prehistory. R-M269 has been found, for instance, at a rate of ~44% among remains dating from the 10th Century at Punta Azul, in the Canary Islands. These remains have been linked to the Bimbache (or Bimape), a subgroup of the Guanche.[42] In living males, it peaks in parts of North Africa, especially Algeria, at a rate of 10%.[43] In Sub-Saharan Africa, R-M269 appears to peak in Namibia, at a rate of 8% among Herero males.[29] (The table below lists in more detail the frequencies of M269 in regions in Asia, Europe, and Africa.)

Apart from undiverged, basal R-M269*, there are (as of 2017) two primary branches of R-M269:

R-L23* and its subclades are by far the most common forms of R-M269. R-PF7558 is now rare, but has been reported anecdotally at low levels throughout Western Eurasia (Europe and the Middle East).[44]

R-L23 (Z2105/Z2103; a.k.a. R1b1a1a2a) has been reported among the peoples of the Idel-Ural (by Trofimova et al. 2015): 21 out of 58 (36.2%) of Burzyansky District Bashkirs, 11 out of 52 (21.2%) of Udmurts, 4 out of 50 (8%) of Komi, 4 out of 59 (6.8%) of Mordvins, 2 out of 53 (3.8%) of Besermyan and 1 out of 43 (2.3%) of Chuvash were R1b-L23.[45]

Subclades within the paragroup R-M269(xL23) – that is, R-M269* and/or R-PF7558 – appear to be found at their highest frequency in the central Balkans, especially Kosovo with 7.9%, Macedonia 5.1% and Serbia 4.4%.[6] Unlike most other areas with significant percentages of R-L23, Kosovo, Poland and the Bashkirs of south-east Bashkortostan are notable in having a high percentage of R-L23(xM412) also known as R1b1a1a2a(xR1b1a1a2a1) – at rates of 11.4% (Kosovo), 2.4% (Poland) and 2.4% south-east Bashkortostan.[6] (This Bashkir population is also notable for its high level of R-M73 (R1b1a1a1), at 23.4%.[6]) Five individuals out of 110 tested in the Ararat Valley of Armenia belonged to R-M269(xL23) and 36 to R-L23*, with none belonging to known subclades of L23.[46]

In 2009, DNA extracted from the femur bones of 6 skeletons in an early-medieval burial place in Ergolding (Bavaria, Germany) dated to around AD 670 yielded the following results: 4 were found to be haplogroup R1b with the closest matches in modern populations of Germany, Ireland and the USA while 2 were in Haplogroup G2a.[47]

The following gives a summary of most of the studies which specifically tested for M269, showing its distribution (as a percentage of total population) in Europe, North Africa, the Middle East and Central Asia as far as China and Nepal.

The phylogeny of R-M269 according to ISOGG 2017:

M269/PF6517


R-M269* (R1b1a1a2*)


L23/PF6534/S141


R-L23* (R1b1a1a2a*)


L51/M412/PF6536/S167


R-L51*/R-M412* (R1b1a1a2a1*)


L151/PF6542


R-L151* (R1b1a1a2a1a*)


M405/U106/S21


R-U106/R-M405/R-S21 (R1b1a1a2a1a1)



P312/PF6547/S116

R-P312 (R1b1a1a2a1a2)


AM01876/S1194

R-S1194 (R1b1a1a2a1a3)


A8051

R-A8051 (R1b1a1a2a1a4)



PF7589/Z2118

R-PF7589 (R1b1a1a2a1b)



CTS1078/Z2103

R-Z2103 (R1b1a1a2a2)



PF7558

R-PF7558 (R1b1a1a2b)



R1b1a1b (R-V1636)

R-V1636 (R1b1a1b) has no known subclades, is rare and has been found mostly in the broader Mediterranean region.

R1b1a2 (R-V88)

R1b1a2 (PF6279/V88; previously R1b1c) is defined by the presence of SNP marker V88, the discovery of which was announced in 2010 by Cruciani et al.[31] Apart from individuals in southern Europe and Western Asia, the majority of R-V88 was found in the Sahel among populations speaking Afroasiatic languages of the Chadic branch:

Region Population Country Language N Total% R1b1c (R-V88) R1b1a1a2 (R-M269) R1b1c* (R-V88*) R1b1c3 (R-V69)
N Africa Composite Morocco AA3380.0%0.3%0.6%0.3%0.0%
N Africa Mozabite Berbers Algeria AA/Berber 673.0%3.0%0.0%3.0%0.0%
N Africa Northern Egyptians Egypt AA/Semitic 496.1%4.1%2.0%4.1%0.0%
N Africa Berbers from Siwa Egypt AA/Berber 9328.0%26.9%1.1%23.7%3.2%
N Africa Baharia Egypt AA/Semitic 417.3%4.9%2.4%0.0%4.9%
N Africa Gurna Oasis Egypt AA/Semitic 340.0%0.0%0.0%0.0%0.0%
N Africa Southern Egyptians Egypt AA/Semitic 695.8%5.8%0.0%2.9%2.9%
C Africa Songhai Niger NS/Songhai 100.0%0.0%0.0%0.0%0.0%
C Africa Fulbe Niger NC/Atlantic 714.3%14.3%0.0%14.3%0.0%
C Africa Tuareg Niger AA/Berber 224.5%4.5%0.0%4.5%0.0%
C Africa Ngambai Chad NS/Sudanic 119.1%9.1%0.0%9.1%0.0%
C Africa Hausa Nigeria (North) AA/Chadic 1020.0%20.0%0.0%20.0%0.0%
C Africa Fulbe Nigeria (North) NC/Atlantic 320.0%0.0%0.0%0.0%0.0%
C Africa Yoruba Nigeria (South) NC/Defoid 214.8%4.8%0.0%4.8%0.0%
C Africa Ouldeme Cameroon (Nth) AA/Chadic 2295.5%95.5%0.0%95.5%0.0%
C Africa Mada Cameroon (Nth) AA/Chadic 1782.4%82.4%0.0%76.5%5.9%
C Africa Mafa Cameroon (Nth) AA/Chadic 887.5%87.5%0.0%25.0%62.5%
C Africa Guiziga Cameroon (Nth) AA/Chadic 977.8%77.8%0.0%22.2%55.6%
C Africa Daba Cameroon (Nth) AA/Chadic 1942.1%42.1%0.0%36.8%5.3%
C Africa Guidar Cameroon (Nth) AA/Chadic 966.7%66.7%0.0%22.2%44.4%
C Africa Massa Cameroon (Nth) AA/Chadic 728.6%28.6%0.0%14.3%14.3%
C Africa Other Chadic Cameroon (Nth) AA/Chadic 475.0%75.0%0.0%25.0%50.0%
C Africa Shuwa Arabs Cameroon (Nth) AA/Semitic 540.0%40.0%0.0%40.0%0.0%
C Africa Kanuri Cameroon (Nth) NS/Saharan 714.3%14.3%0.0%14.3%0.0%
C Africa Fulbe Cameroon (Nth) NC/Atlantic 1811.1%11.1%0.0%5.6%5.6%
C Africa Moundang Cameroon (Nth) NC/Adamawa 2166.7%66.7%0.0%14.3%52.4%
C Africa Fali Cameroon (Nth) NC/Adamawa 4820.8%20.8%0.0%10.4%10.4%
C Africa Tali Cameroon (Nth) NC/Adamawa 229.1%9.1%0.0%4.5%4.5%
C Africa Mboum Cameroon (Nth) NC/Adamawa 90.0%0.0%0.0%0.0%0.0%
C Africa Composite Cameroon (Sth) NC/Bantu 900.0%1.1%0.0%1.1%0.0%
C Africa Biaka Pygmies CAR NC/Bantu 330.0%0.0%0.0%0.0%0.0%
W Africa Composite — 1230.0%0.0%0.0%0.0%0.0%
E Africa Composite — 4420.0%0.0%0.0%0.0%0.0%
S Africa Composite — 1050.0%0.0%0.0%0.0%0.0%
TOTAL1822

V88

un-defined

R-V88* (R1b1c*)


M18

R-M18 (R1b1c1)


V35

R-V35 (R1b1c2)


V69

R-V69 (R1b1c3)



As can be seen in the above data table, R1b1c is found in northern Cameroon in west central Africa at a very high frequency, where it is considered to be caused by a pre-Islamic movement of people from Eurasia.[29][48]

R1b1a2a (R-M18)

R1b1a2a is a sub-clade of R-V88, which is defined by the presence of SNP marker M18.[7] It has been found only at low frequencies in samples from Sardinia[40][49] and Lebanon.[50]

R1b1b (R-PH155)

The other secondary branch of R1b is R-PH155 (R1b1b), which is extremely rare and defined by the presence of PH155.[51] ISOGG (2017) cites two primary branches: R-M335 (R1b1b1) and R-PH200 (R1b1b2).

Another defining SNP of R1b1b, M335, was first documented in 2004, when an example was discovered in Turkey, which was classified at that time as R1b4.[21]

Other examples of R-PH155 have been found in Bahrain, Bhutan, Tajikistan and Xinjiang.

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
L     T [χ 6] K2a [χ 7] K2b [χ 8]   K2c   K2d  K2e [χ 9]  
K2a1                    K2b1 [χ 10]    P [χ 11]
NO    S [χ 12]  M [χ 13]    P1     P2
NO1    Q   R
N O
  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. PMID 24166809. doi:10.1002/humu.22468.
  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 A0'1'2'3'4.
  4. ↑ Haplogroup A1 is also known as A1'2'3'4.
  5. ↑ Haplogroup LT (L298/P326) is also known as Haplogroup K1.
  6. ↑ 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.
  7. ↑ Haplogroup K2a (M2308) and the new subclade K2a1 (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. PMC 4884158 Freely accessible. PMID 27111036. doi:10.1038/ng.3559. In the past, other haplogroups, including NO1 (M214) and K2e had also been identified with the name "K2a".
  8. ↑ Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS.
  9. ↑ Haplogroup K2e (K-M147) was previously known as "Haplogroup X" and "K2a" (but is a sibling subclade of the present K2a).
  10. ↑ Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure.
  11. ↑ Haplogroup P (P295) is also klnown as K2b2.
  12. ↑ Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)
  13. ↑ Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)

References

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  3. ↑ Massive migration from the steppe is a source for Indo-European languages in Europe, Haak et al, 2015
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  5. ↑ Eight thousand years of natural selection in Europe, Mathieson et al, 2015
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  39. ↑ R1b1a1a2 (R-M269) was previously R1b1a2, From 2003 to 2005, what is now R1b1a2 was designated R1b3. From 2005 to 2008, it was R1b1c. From 2008 to 2011, it was R1b1b2.
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  52. ↑ About R1a and R1b from Ural epic story. Artem Lukichev (c)
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Unannotated references

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