Haplogroup N-M231

Haplogroup N (M231) is a Y-chromosome DNA haplogroup defined by the presence of the single-nucleotide polymorphism (SNP) marker M231.[Phylogenetics 1]

This article is about the human Y-DNA haplogroup. For the human mtDNA haplogroup, see Haplogroup N (mtDNA).
Haplogroup N
Possible time of origin36,800 [95% CI 34,300–39,300] years before present (YFull[1])

44,700 or 38,300 ybp depending on mutation rate[2]

41,900 [95% CI 40,175-43,591] ybp[3]
Coalescence age21,700 [95% CI 19,500–23,900] ybp (YFull[1])

25,313 [95% CI 21,722-28,956] ybp[3]
Possible place of originEast Asia[4][5]
AncestorNO
Defining mutationsM231
Highest frequenciesNganassan 58%[3]-94.1%,[6] Yakuts 81.8%[7]-94.6%[8] (Xu 2015), Ugrians 77.8%[3] (Khanty 64.3%[9]-89.3%,[10] Mansi 76%[10]), Udmurts 77.8%,[3] Komi 33.3%[9]-79.5%,[6] Nenets 75%-92.9%[3] (Tundra Nenets 97.9%,[6] Forest Nenets 98.8%[6]), Finns 51-61% (Purps 2014), Tuvans 27.2-54.5% (Kharkov 2013), Nanai 46.2%[11][12][3] (20% Hezhe in the PRC,[11] 44.6% Nanai in Russia,[3] 83.8% members of the Samar clan in the Gorin area of the Khabarovsk Territory[12]), Arkhangelsk Russians 42.6% (Arkhangelsk 44.3%,[3] Pinega 40.8%[3]), Mari 41.2%,[3] Estonian 30.6%[3]-40%(Purps 2014), Saami 40%, Chuvash 36%,[3] Buryats 34.5% (20.2%,[13] 25.0%,[14] 30.9%,[15] 48.0%[16]), Koryaks 33.3%,[6] Latvian 30%(Purps 2014)-43.0%,[3] Volga Tatars 27.8%,[3] Siberian Tatars 27.6%,[3] Lithuanian 25%(Purps 2014)-40.5%,[3] Teleuts 25.0%,[6] Northern Altaians 21.8% (18.0%[17][9]-24.6%[18]), Pskov Russians 22.7%,[3] Swedish 9-22% (Purps 2014), Bashkirs 17.3%,[3] Sibe 17.1%[11]-18.0%,[19] Mordvins 12.5% (10%[3] - 13.3%[3]), Mongols 11%,[20][15][11][14][21][22] Kalmyks 10.4% (Torguud 3.4%, Derbet 5.1%, Buzava 5.3%, Khoshut 38.2%),[23][22] Manchus 10% (5.8%,[15] 9.1%,[19] 11.6%,[19] 12.5%,[19] 14.3%[11]), Belarusians 9.7%,[3] Central-Southern Russians 9.1% (Tver 13.2%,[24] Kursk 12.5%[24]-13.3%,[3] Belgorod 11.9%,[3] Kostroma 11.8%,[3] Smolensk 7.0%,[3] Voronezh 6.3%,[3] Oryol 5.5%[3]), Ukrainians 9.0%,[3] Southern Altaians 7.1% (4.2%[18]-9.7%[9]), Han Chinese 6.77% (0% to 21.4%),[19] Ulchi 5.8%,[25] Tibetans 5.65%,[26] Kazakhs 5.33%[27] (Suan 0%, Qangly 0%, Oshaqty 0%, Jetyru 1.2%, Naiman 1.3%, Dulat 1.6%, Argyn 2.0%, Alimuly 2.5%, Ysty 3.5%, Kerey 3.6%, Baiuly 3.9%, Alban 4.3%, Qongyrat 7.4%, Qypshaq 10.3%, Jalair 10.9%, Qozha 16.7%, Syrgeli 65.6%), Uyghurs 4.89% (2.8%,[28] 4.8%,[19] 4.99%,[29] 6.0%,[15] 8.6%[11]), Koreans 4.41% (1.8% Seoul-Gyeonggi,[14] 3.0% Daejeon,[30] 4.0% Seoul,[30] 4.2% Chungcheong,[14] 4.4% Jeolla,[14] 4.8% Gyeongsang,[14] 6.3% Gangwon,[14] 6.58% Korean Genome Project (mostly from Ulsan),[31] 6.9% Jeju[14]), Japanese 1.9% (0%,[2] 0.8%,[32] 0.9%,[33] 1.7%,[34] 2.5%,[14] 4.3%,[35] 4.8%,[15] 6.4%[11])

It is most commonly found in males originating from northern Eurasia. It also has been observed at lower frequencies in populations native to other regions, including the Balkans, Central Asia and East Asia

Origins
Estimated prehistoric migration routes for Y-chromosome haplogroup N lineage.[36]

Haplogroup NO-M214 – its most recent common ancestor with its sibling, haplogroup O-M175 – is estimated to have existed about 36,800–44,700 years ago.[1][37][2]

It is generally considered that N-M231 arose in East Asia approximately 19,400 (±4,800) years ago and re-populated northern Eurasia after the Last Glacial Maximum. Males carrying the marker apparently moved northwards as the climate warmed in the Holocene, migrating in a counter-clockwise path (through modern China and Mongolia), to eventually become concentrated in areas as far away as Fennoscandia and the Baltic (Rootsi et al. 2006). The apparent dearth of haplogroup N-M231 amongst Native American peoples indicates that it spread after Beringia was submerged (Chiaroni, Underhill & Cavalli-Sforza 2009), about 11,000 years ago.

Distribution
Projected distributions of haplogroup N sub-haplogroups.[36] (A) N*-M231, (B) N1*-LLY22g, (C) N1a-M128, (D) N1b-P43, (E) N1c-M46.

Haplogroup N has a wide geographic distribution throughout northern Eurasia, and it also has been observed occasionally in other areas, including Central Asia and the Balkans.

It has been found with greatest frequency among indigenous peoples of Russia, including Finnic peoples, Mari, Udmurt, Komi, Khanty, Mansi, Nenets, Nganasans, Turkic peoples (Yakuts, Dolgans, Khakasses, Tuvans, Tatars, Chuvashes, etc.), Buryats, Tungusic peoples (Evenks, Evens, Negidals, Nanais, etc.), Yukaghirs, Luoravetlans (Chukchis, Koryaks), and Siberian Eskimos, but certain subclades are very common in Finland, Estonia, Latvia, and Lithuania, and other subclades are found at low frequency in China (Yi, Naxi, Lhoba, Han Chinese, etc.).[38] Especially in ethnic Finnic peoples and Baltic-speaking peoples of northern Europe, the Ob-Ugric-speaking and Northern Samoyed peoples of western Siberia, and Turkic-speaking peoples of Russia (especially Yakuts (McDonald 2005), but also Altaians, Shors, Khakas, Chuvashes, Tatars, and Bashkirs). Nearly all members of haplogroup N among these populations of northern Eurasia belong to subclades of either haplogroup N-Tat or haplogroup N-P43.

Y-chromosomes belonging to N1b-F2930/M1881/V3743, or N1*-CTS11499/L735/M2291(xN1a-F1206/M2013/S11466), have been found in China and sporadically throughout other parts of Eurasia.

N2-Y6503, the other primary subclade of haplogroup N, is extremely rare and is mainly represented among extant humans by a recently formed subclade that is virtually restricted to the countries making up the former Yugoslavia (Bosnia-Herzegovina, Croatia, Serbia, and Montenegro), Hungary and Austria. Other members of N2-Y6503 include a Hungarian with recent ancestry from Suceava in Bukovina, a Slovakian, a few British individuals, and an Altaian.[1]

N* (M231)

Y-chromosomes that display the M231 mutation that defines Haplogroup N-M231, but do not display the CTS11499, L735, M2291 mutations that define Haplogroup N1 are said to belong to paragroup N-M231*.[4]

N-M231* has been found at low levels in China.[4] Out of a sample of 165 Han males from China, two individuals (1.2%) were found to belong to N*.(Karafet et al. 2010).[Footnote 1] One originated from Guangzhou and one from Xi'an.

N1 (CTS11499, Z4762, CTS3750)

In 2014, there was a major change in the definition of subclade N1, when LLY22g was retired as the main defining SNP for N1 because of reports of LLY22g's unreliability. According to ISOGG, LLY22g is problematic because it is a "palindromic marker and can easily be misinterpreted."[4] Since then, the name N1 has been applied to a clade marked by a great number of SNPs, including CTS11499, Z4762, and CTS3750. N1 is the most recent common ancestor of all extant members of Haplogroup N-M231 except members of the rare N2-Y6503 (N2-B482) subclade. The TMRCA of N1 is estimated to be 18,000 years before present (16,300–19,700 BP; 95% CI).[1] The modern populations with the greatest proportions of N-CTS11499 (or N-Z4762) are concentrated in China.

Since the revision of 2014, the position of many examples of "N1-LLY22g" within haplogroup N have become unclear. N-LLY22g has been reported to reach a frequency of up to 30% (13/43) among the Yi people of Butuo County, Sichuan in Southwest China (Hammer et al. 2005, Karafet et al. 2001, and Wen2004b). It is also found in 34.6% of Lhoba people (Wen 2004, Bo Wen 2004).[39] N1-LLY22g* has been found in samples of Han Chinese, but with widely varying frequency:

Other populations in which representatives of N1*-LLY22g have been found include:

N1(xN1a,N1c) was found in ancient bones of Liao civilization:[40]

  • Niuheliang (Hongshan Culture, 6500–5000 BP) 66.7%(=4/6)
  • Halahaigou (Xiaoheyan Culture, 5000–4200 BP) 100.0%(=12/12)
  • Dadianzi (Lower Xiajiadian culture, 4200–3600 BP) 60.0%(=3/5)

N1a (F1206/M2013/S11466)

The N1a2-F1008/L666 clade and N1a1-M46/Page70/Tat are estimated to share a most recent common ancestor in N1a-F1206/M2013/S11466 approximately 15,900 [95% CI 13,900 <-> 17,900] years before present[1] or 17,621 [95% CI 14,952 <-> 20,282] years before present.[3]

N1a1 (M46/Page70/Tat, L395/M2080)

All M46 in Yfull database are M178, being a quarter younger than separation from F1139.[41]

The mutations that define the subclade N-M46[Phylogenetics 2] are M46/Tat and P105. This is the most frequent subclade of N. It arose probably in the region of present-day China, and subsequently experienced serial bottlenecks in Siberia and secondary expansions in eastern Europe (Rootsi et al. 2006). Haplogroup N-M46 is approximately 14,000 years old.

In Siberia, haplogroup N-M46 reaches a maximum frequency of approximately 90% among the Yakuts, a Turkic people who live mainly in the Sakha (Yakutia) Republic. However, N-M46 is present with much lower frequency among many of the Yakuts' neighbors, such as Evenks and Evens.[8] It also has been detected in 5.9% (3/51) of a sample of Hmong Daw from Laos (Cai 2011), 2.4% (2/85) of a sample from Seoul, South Korea (Katoh 2004), and in 1.4% (1/70) of a sample from Tokushima, Japan (Hammer et al. 2005).

The haplogroup N-M46 has a low diversity among Yakuts suggestive of a population bottleneck or founder effect (Pakendorf 2002). This was confirmed by a study of ancient DNA which traced the origins of the male Yakut lineages to a small group of horse-riders from the Cis-Baikal area (Crubézy 2010).

N1a1a (M178)

The subclade N-M178[Phylogenetics 3] is defined by the presence of markers M178 and P298. N-M178* has higher average frequency in Northern Europe than in Siberia, reaching frequencies of approximately 60% among Finns and approximately 40% among Latvians, Lithuanians & 35% among Estonians (Derenko 2007 and Lappalainen 2008).

Miroslava Derenko and her colleagues noted that there are two subclusters within this haplogroup, both present in Siberia and Northern Europe, with different histories. The one that they labelled N3a1 first expanded in south Siberia and spread into Northern Europe. Meanwhile, the younger subcluster, which they labelled N3a2, originated in south Siberia (probably in the Baikal region)(Derenko 2007).

N-M178 was also found in two Na-Dené speaking Tłı̨chǫs in North America.[42]

N1a2 (F1008/L666)

N1a2a-M128 and N1a2b-B523/P43 are estimated to share a most recent common ancestor in N1a2-F1008/L666 approximately 8,600 [95% CI 7,500 <-> 9,800] years before present[1] or 9,314 [95% CI 7,419 <-> 11,264] years before present.[3]

At least three of six tested male specimens from the "Early Neolithic" (ceramic-using hunter-gatherer of approximately 7200–6200 years before present) layer at the Shamanka archaeological site near the southern end of Lake Baikal have been found to belong to N1a2-L666.[43]

N1a2a-M128
Haplogroup N-M128
Possible place of originAsia
AncestorN1c2 (F1008/L666)
Defining mutationsM128

This subclade is defined by the presence of the marker M128.[Phylogenetics 4] N-M128 was first identified in a sample from Japan (1/23 = 4.3%) and in a sample from Central Asia and Siberia (1/184 = 0.5%) in a preliminary survey of worldwide Y-DNA variation.[35] Subsequently, it has been found with low frequency in some samples of the Manchu people, Sibe people, Evenks, Koreans, Han Chinese, Hui, Tibetans, Vietnamese, Bouyei people, Kazakhs, Uzbeks, Uyghurs, Salars, Tu, Mongols, the Buzava tribe of Kalmyks,[23] Khakas, and Komis.[44]

A number of Han Chinese, an Ooled Mongol, a Qiang, and a Tibetan were found to belong to a sister branch (or branches) of N-M128 under paragroup N-F1154*.[45]

N1a2b (P43)

Haplogroup N-P43[Phylogenetics 5] is defined by the presence of the marker P43. It has been estimated to be about five thousand years old (TMRCA 4,700 [95% CI 3,800 <-> 5,600] ybp[1] or 4,727 [95% CI 3,824 <-> 5,693] years before present[3]). It has been found very frequently among Northern Samoyedic peoples and speakers of Ob-Ugric languages, and it also has been observed with low to moderate frequency among speakers of some other Uralic languages, Turkic peoples, Mongolic peoples, Tungusic peoples, and Siberian Yupik people.

The highest frequencies of N-P43 are observed among north-west Siberian populations: 92% (35/38)[46] in a sample of Nganasan, 78% (7/9)[47][48] in a sample of Enets, 78% (21/27)[24] in a sample of Khants, 75% (44/59)[46] in a sample of Tundra Nenets, 69% (29/42)[3] in another sample of Nenets, 60% (15/25)[10] in a sample of Mansi, 57% (64/112)[9] in another sample of Khants, 54% (27/50)[3] in another sample of Nganasan, 45% (40/89)[46] in a sample of Forest Nenets, 38% (18/47)[49] in a third sample of Khants, and 25% (7/28)[10] in a fourth sample of Khants. In Europe, the N-P43 types have their highest frequency of 20% among Volga-Uralic populations. The extreme western border of the spread of N-P43 is Finland, where this haplogroup occurs only at marginal frequency – 0.4%. Yet N-P43 is quite frequent among Vepsas (17.9%), a small Finnic population living in immediate proximity to Finns, Karelians and Estonians.[50]

Haplogroup N-P43 forms two distinctive subclusters of STR haplotypes, Asian and European, the latter mostly distributed among Finno-Ugric-speaking peoples and related populations (Rootsi et al. 2006).

N1a2b1-B478

The TMRCA of N-B478 has been estimated to be 3,007 [95% CI 2,171 <-> 3,970] years before present.[3] It is one of the most prevalent Y-DNA haplogroups among indigenous populations of northwestern Siberia: 69.0% (29/42) Nenets, 50.0% (25/50) Nganasan, 22.2% (12/54) Dolgan from Taymyr, 7.0% (3/43) Selkup, 1.6% (1/63) Ob-Ugrian. It is also quite prevalent among populations of Central Siberia, Southern Siberia, Mongolia, and Kyrgyzstan: 17.9% (17/95) Tuvan, 15.5% (27/174) Khakas, 13.0% (6/46) Tozhu Tuvans,[22] 8.7% (2/23) Shor, 8.3% (2/24) Even, 8.2% (5/61) Altaian, 5.3% (3/57) Evenk, 5.0% (19/381) Mongol, 4.9% (3/61) Sart-Kalmak (Kyrgyzstan),[22] 4.2% (9/216) Yakut, 2.1% (1/47) Torgut (Mongolia),[22] 1.4% (1/69) Derbet (Kalmykia),[22] 0.9% (1/111) Buryat. A geographically outlying member has been found in a sample of Chuvash (1/114 = 0.88%).[3]

Karafet et al. (2018) observed N-P63, which appears to be roughly phylogenetically equivalent to N-B478, in 91.2% (31/34) Nganasan, 63.8% (30/47) Tundra Nenets, 42.7% (35/82) Forest Nenets, 14.0% (8/57) Dolgan, 7.0% (9/129) Selkup, 3.3% (3/91) Evenk, 2.7% (2/75) Mongol, 2.6% (2/78) Komi, 2.5% (2/80) Buryat, and 2.0% (2/98) Altai Kizhi.[6] This haplogroup was not observed in samples of Yukaghir (0/10), Koryak (0/11), Teleut (0/40), Ket (0/44), Yakut (0/62), or Khanty (0/165) populations.[6]

N1b (F2930)

Haplogroup N1b has been predominantly found in the Yi people, a Tibeto-Burman speaking ethnic group in southwestern China who originated from ancient Qiang tribes in northwestern China.[51] However, it also has been found in people all over China and in some samples in Poland, Belarus, Russia, Mongolia, Korea, Japan and Vietnam

N2 (Y6503)

N2 (Y6503/FGC28528; B482/FGC28394/Y6584) – a primary branch of haplogroup N-M231, is now represented mainly by a subclade, N-FGC28435, that has spread probably some time in the first half of the second millennium CE[52] and that has been found in individuals from Serbia, Croatia, Bosnia and Herzegovina, Montenegro, and Turkey (Istanbul).[53][52]

N-Y7310 (or N-F14667) subsumes N-FGC28435 and likewise probably descends from a common ancestor who has lived some time in the first half of the last millennium. However, members of N-Y7310(xFGC28435) exhibit a greater geographic range, including an individual from Rostov Oblast of Russia and a Hungarian individual with ancestry from Suceava, Bukovina.[1]

Other branches of N-P189 include members from Italy, Romania, Slovakia, and England (Devon[1]).[54] The most recent common ancestor of all the aforementioned extant N-P189 lineages dates back to some time in the second, third, or even as early as the latter half of the fourth millennium BCE.[1] An archaeological specimen attributed to the Botai culture of northern Kazakhstan of the fourth millennium BCE may belong to a pre-N-P189 branch.[55]

Lineages that belong to N-Y6503(xP189) and are only distantly related (with a time to most recent common ancestor estimated to be greater than 10,000 years before present[1]) to the aforementioned members of N-P189 have been found in an individual from the present-day Altai Republic[1] and probably also in an archaeological specimen attributed to the Iron Age Mezőcsát culture of what is now Hungary (approx. 2,900 years before present)[56] and in an archaeological specimen attributed to the Kitoi culture of ceramic-using foragers of the area around Lake Baikal (approx. 6,700 years before present).[55]

Ancient peoples

Most samples from the Liao civilization in northeastern China and northern Korea belonged to y-DNA N. N has been found in many samples of Neolithic human remains exhumed from northeastern China and the circum-Baikal area of southern Siberia. It is thus suggested that the ancestors of the Uralic peoples and of the Turkic-Yakut peoples may have originated in this region about 8000–6000 years ago.[57]

Phylogeny

Phylogenetic tree

In the following tree the nomenclature of three sources is separated by slashes: ISOGG Tree 10 December 2017 (ver.12.317)

Phylogenetic history
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, the major research groups came together and formed the Y-Chromosome Consortium (YCC). They published a joint paper that created a single new tree that all agreed to use. Later, a group of citizen scientists with an interest in population genetics and genetic genealogy formed a working group to create an amateur tree aiming at being above all timely. 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 2006 ISOGG 2007 ISOGG 2008 ISOGG 2009 ISOGG 2010 ISOGG 2011 ISOGG 2012
N-LLY22g12VIII1U25Eu16H5FN*NN1N1-------
N-M12812VIII1U25Eu16H5FN1N1N1aN1a-------
N-P6312VIII1U25Eu16H5FN2N2aN1b1N1b1-------
N-TAT12VIII1I26Eu13H5FN3*N3N1cN1c-------
N-M17816VIII1I26Eu14H5FN3a*M178N1c1N1c1-------
N-P2116VIII1I26Eu14H5FN3a1N3a1N1c1aN1c1a-------

Sources The following research teams per their publications were represented in the creation of the YCC Tree.

Unreliable mutations (SNPs and UEPs)

The b2/b3 deletion in the AZFc region of the Y-chromosome appears to have occurred independently on at least four different occasions. Therefore, this deletion should not be taken as a unique event polymorphism defining this branch of the Y-chromosome tree (ISOGG 2012).

Genetics

Y-DNA N subclades
  • N-M231

Y-DNA backbone tree
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

Footnotes
  1. In Karafet et al. 2010

Work cited
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  3. Ilumäe 2016
  4. ISOGG, 2016, Y-DNA Haplogroup N and its Subclades – 2016 22 August 2016).
  5. (Rootsi et al. 2006)
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Journals

Websites

Further reading

Phylogenetics
  1. The b2/b3 deletion in the AZFc region of the human Y-chromosome is a characteristic of Haplogroup N-M231 haplotypes. This deletion, however, appears to have occurred independently on four different occasions. Therefore this deletion should not be thought as a unique event polymorphism contributing to the definition of this branch of the Y-chromosome tree (ISOGG 2012).
  2. This table shows historic names for N-M46 (AKA N-Tat) from peer reviewed literature.
    YCC 2002/2008 (Shorthand) N-M46/N-TAT
    Jobling and Tyler-Smith 200012
    Underhill 2000VIII
    Hammer 20011I
    Karafet 200126
    Semino 2000Eu13
    Su 1999H5
    Capelli 2001F
    YCC 2002 (Longhand)N3*
    YCC 2005 (Longhand)N3
    YCC 2008 (Longhand)N1c
    YCC 2010r (Longhand)N1c
  3. This table shows historic names for N-M178 from peer reviewed literature.
    YCC 2002/2008 (Shorthand) N-M178
    Jobling and Tyler-Smith 200016
    Underhill 2000VIII
    Hammer 20011I
    Karafet 200126
    Semino 2000Eu14
    Su 1999H5
    Capelli 2001F
    YCC 2002 (Longhand)N3a*
    YCC 2005 (Longhand)M178
    YCC 2008 (Longhand)N1c1
    YCC 2010r (Longhand)N1c1
  4. This table shows historic names for N-M128 from peer reviewed literature.
    YCC 2002/2008 (Shorthand) N-M128
    Jobling and Tyler-Smith 200012
    Underhill 2000VIII
    Hammer 20011U
    Karafet 200125
    Semino 2000Eu16
    Su 1999H5
    Capelli 2001F
    YCC 2002 (Longhand)N1
    YCC 2005 (Longhand)N1
    YCC 2008 (Longhand)N1a
    YCC 2010r (Longhand)N1a
  5. This branch is sometimes called N1b in early trees.
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