Haplogroup O-M119

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Haplogroup O-M119
Possible time of origin 34,100 or 29,200 years ago[1]

30,100 [95% CI 27,800 <-> 32,400] years before present[2]
Possible place of origin Southeast Asia or Southern China
Coalescence age 15,000 [95% CI 13,200 <-> 16,900] years before present[2]
Ancestor O-M175 > O-F265
Defining mutations M119

In human genetics, Haplogroup O-M119 is a Y-chromosome DNA haplogroup. Haplogroup O-M119 is a descendant branch of O-F265, one of two extant primary subclades of Haplogroup O-M175. The same clade previously has been labeled as O-MSY2.2.[3]


The Haplogroup O-M119 branch is believed to have evolved during the Late Pleistocene (Upper Paleolithic) in Southeast Asia.[citation needed]

Paleolithic migrations[edit]

A 2010 study by Karafet suggests haplogroup O-M119 was part of a four-phase colonization model in which Paleolithic migrations of hunter-gatherers shaped the primary structure of current Y-Chromosome diversity of Maritime Southeast Asia. Neolithic incursions made only a minor impact on the paternal gene pool, despite the large cultural impact of the Austronesian expansion. Approximately 5000 BCE, Haplogroup O-M119 coalesced at Sundaland and migrated northwards to as far as Taiwan, where O-M50 constitutes some 90% of the Aboriginal Y-DNA, being the main haplogroup that can be directly linked to the Austronesian expansion in phase 3 (Karafet 2010).

Taiwan homeland[edit]

A study by Li in 2008 concluded that in contrast to the Taiwan homeland hypothesis, Island Southeast Asians do not have a Taiwan origin based on their paternal lineages. According to their results, lineages within Maritime Southeast Asia did not originate from Taiwanese aborigines as linguistic studies suggest. Taiwan aborigines and Indonesians were likely to have been derived from the Tai–Kadai-speaking populations based on their paternal lineages, and thereafter evolved independently of each other (Li 2008).

The strongest positive correlation between Haplogroup O-M119 and ethno-linguistic affiliation is that which is observed between this haplogroup and the Austronesians. The peak frequency of Haplogroup O-M119 is found among the aborigines of Taiwan, precisely the region from which linguists have hypothesized that the Austronesian language family originated. A slightly weaker correlation is observed between Haplogroup O-M119 and the Han Chinese populations of southern China, as well as between this haplogroup and the Tai–Kadai-speaking populations of southern China and Southeast Asia. The distribution of Tai–Kadai languages in Thailand and other parts of Southeast Asia outside of China has long been believed, for reasons of traditional linguistic geography, to reflect a recent invasion of Southeast Asia by Tai–Kadai-speaking populations originating from southeastern China, and the somewhat elevated frequency of Haplogroup O-M119 among the Tai–Kadai populations, coupled with a high frequency of Haplogroup O-M95, which is a genetic characteristic of the Austroasiatic-speaking peoples of Southeast Asia, suggests that the genetic signature of the Tai–Kadai peoples' affinity with populations of southeastern China has been weakened due to extensive assimilation of the earlier Austroasiatic residents of the lands which the Tai–Kadai peoples invaded. Also, it has been noted that Haplogroup O-M119 lineages among populations of continental Southeast Asia outside of China display a reduced level of diversity when compared with populations of South China and insular Southeast Asia, which may be evidence of a bottleneck associated with the westward migration and settlement of ancestral Tai–Kadai-speaking populations in Indochina.[citation needed]


Haplogroup O-M119 lineages are found primarily in Southeast Asian populations of Malaysia, Vietnam, Indonesia, the Philippines, southern China and Taiwan (ISOGG 2010). High frequencies of this haplogroup have been found in populations spread in an arc through southeastern China, Taiwan, the Philippines, and Indonesia. It has been found with generally lower frequency in samples from Oceania, mainland Southeast Asia, Southwest China, Northwest China, North China, Northeast China, Korea, Japan, North Asia, and Central Asia.

Population Percentage Count Source SNPs
Nias 100.0% 60 Karafet 2010 M119
Nias 99.8% 407 vanOven 2011 M119
Taiwanese aborigines 89.6% 48 Karafet 2010 M119
Mentawai 86.5% 74 Karafet 2010 M119
Aboriginal Taiwanese 83.4% 223 Tajima 2004 M119
Taiwan (aborigines) 71.8% 39 Hurles 2005 M119(xM101)
Taiwan (aborigines) 68.9% 74 Underhill 2000 M119(xM101)
Hlai/Cun 58% - Li 2008 -
Tagalog (Philippine subgroup) 46.0% 50 Tajima 2004 M119
Philippines 35.7% 28 Hurles 2005 M119(xM101)
Kota Kinabalu 29.2% 65 Hurles 2005 M119
Trobriand Islands 28% - Kayser 2003 -
Li (Hlai) 26.5% 34 Xue 2006 M119
Malay (near Kuala Lumpur) 25.0% 12 Tajima 2004 M119
Han (East China) 24.0% 167 Yan 2011 M119
Han Chinese 23% - Kayser 2003 -
Java 23% - Kayser 2003 -
Nusa Tenggara 23% - Kayser 2003 -
Banjarmasin 22.7% 22 Hurles 2005 M119(xM101)
Balinese 21.1% 641 Karafet 2010 M119
Mandar 20.4% 54 Karafet 2010 M119(xP203)
Tujia 20% - Su 1999 -
Han (Meixian) 20.0% 35 Xue 2006 M119
Buka 20.0% 10 Scheinfeldt 2006 M119
Thin Board Mien 18.2% 11 Cai 2011 M119(xM110)
Majuro (Marshall Islands) 18.2% 11 Hurles 2005 M50
Admiralty Islands 18% - Kayser 2008 -
Balinese 18% - Karafet 2005 -
Sui 18% - Xie 2004 -
Zhuang 18% - Su 1999 -
Malagasy 17.1% 35 Hurles 2005 M50
Han (South China) 16.9% 65 Yan 2011 M119
Qiang 15.2% 33 Xue 2006 M119
Borneo 15% - Kayser 2003 -
Han Chinese 15% - Tajima 2004 -
Java 14.8% 61 Karafet 2010 M119
She 14.7% 34 Xue 2006 M119
Han (Chengdu) 14.7% 34 Xue 2006 M119
Manus 14.3% 7 Scheinfeldt 2006 M119
Palyu 13.3% 30 Cai 2011 M119
Batak Toba 13.2% 38 Karafet 2010 M119(xM110)
Sumba 12.6% 350 Karafet 2010 M119
Micronesia 12.5% 16 Karafet 2010 M119(xP203, M110)
Guizhou Miao 12.2% 49 Cai 2011 M119(xM110)
Lowland Kimmun 12.2% 41 Cai 2011 M119(xM110)
Thai (Northern Thailand) 11.8% 17 He 2012 P203(xM101)
Bunu 11.1% 36 Cai 2011 M119(xM110)
Filipinos 10.4% 48 Karafet 2010 M119
Zhuang 10% - Hammer 2006 -
Mountain Straggler Mien 10.0% 20 Cai 2011 M119(xM110)
Han (China & Taiwan) 9.7% 165 Karafet 2010 M119(xM110)
Flores 9.6% 394 Karafet 2010 M119(xM110)
Zhuang 9.6% 166 Chen 2006 -
Han (Taiwan) 9.5% 21 Tajima 2004 M119
Han (Yili) 9.4% 32 Xue 2006 M119
Borneo (Indonesia) 9.3% 86 Karafet 2010 M119
Bougainville 9.2% 65 Scheinfeldt 2006 M119
Top Board Mien 9.1% 11 Cai 2011 M119(xM110)
Northern Mien 9.1% 33 Cai 2011 M119(xM110)
Northern She 8.9% 56 Cai 2011 M119(xM110)
Thai (Chiang Mai & Khon Kaen) 8.8% 34 Tajima 2004 M119
Hui 8.6% 35 Xue 2006 M119
Mosuo (Ninglang, Yunnan) 8.5% 47 Wen 2004 M119(xM110)
Tonga 8.3% 12 Karafet 2010 M119(xP203, M110)
Tujia (Jishou, Hunan) 8.2% 49 Karafet 2010 P203
Hlai/Cun 8% - Li 2008 -
Ewenki (China) 7.7% 26 Xue 2006 M119
Xibe 7.3% 41 Xue 2006 M119
Hunan Miao 7.0% 100 Cai 2011 M119(xM110)
Tujia 7% - Xie 2004 -
Miao (China) 6.9% 58 Karafet 2010 P203
Katu 6.7% 45 Cai 2011 M119(xM110)
Moluccas 6.7% 30 Karafet 2010 M119
Han (Lanzhou) 6.7% 30 Xue 2006 M119
Kinh 6.7% 15 Cai 2011 M119(xM110)
Kinh (Hanoi, Vietnam) 6.6% 76 He 2012 P203(xM101)
Southern Mien 6.5% 31 Cai 2011 M119(xM110)
Malaysia 6.3% 32 Karafet 2010 M119(xM110)
Yunnan Miao 6.1% 49 Cai 2011 M119(xM110)
Northern Han 6.1% 49 Tajima 2004 M119
Comorians 6.0% 381 Msaidie 2010 M50=22
Bai (Dali, Yunnan) 6.0% 50 Wen 2004 M119(xM110)
Kyrgyz (Kyrgyzstan) 5.8% 52 Wells 2001 M119
Vietnam 5.7% 70 Karafet 2010 P203
Yao (Liannan, Guangdong) 5.7% 35 Xue 2006 M119
Samoa 5.6% 18 Karafet 2010 P203
Daur 5.1% 39 Xue 2006 M119
Cham (Binh Thuan, Vietnam) 5.1% 59 He 2012 M119(xM50)
Dungan (Kyrgyzstan) 5.0% 40 Wells 2001 M119
Maewo (Vanuatu) 4.5% 44 Karafet 2010 M119(xP203)
Korean 4.4% 45 Wells 2001 M119
Western Mien 4.3% 47 Cai 2011 M119(xM110)
Pumi (Ninglang, Yunnan) 4.3% 47 Wen 2004 M119(xM110)
Mongolian 4.2% 24 Wells 2001 M119
Western Samoa 4.0% 25 Hurles 2005 M119(xM101, M50)
New Ireland 3.7% 109 Scheinfeldt 2006 M119
Bo 3.6% 28 Cai 2011 M119(xM110)
Japanese 3.4% 263 Nonaka 2007 M119(xM101, M50)
Lembata 3.3% 92 Karafet 2010 M119(xM110)
Korean 3.2% 216 Kim 2007 M119
Han (North China) 3.1% 129 Yan 2011 M119(xM110)
Papua New Guinea Highlands 3.0% 33 Karafet 2010 P203
Manchu 2.9% 35 Xue 2006 M119
Han (Harbin) 2.9% 35 Xue 2006 M119
Buyi 2.9% 35 Xue 2006 M119
Yao (Bama, Guangxi) 2.9% 35 Xue 2006 M119
West New Britain 2.8% 249 Scheinfeldt 2006 M119
Lavongai 2.3% 43 Scheinfeldt 2006 M119
Laven 2.0% 50 Cai 2011 M119(xM110)
Yi (Shuangbai, Yunnan) 2.0% 50 Wen 2004 M119(xM110)
Hmong Daw (Laos) 2.0% 51 Cai 2011 M119(xM110)
She 2.0% 51 Karafet 2010 P203
Japanese (Kyushu) 1.9% 104 Tajima 2004 M119
Vanuatu 1.9% 52 Hurles 2005 M50
Yao (Guangxi) 1.7% 60 Karafet 2010 P203
Uygur 1.4% 70 Xue 2006 M119
East New Britain 1.4% 145 Scheinfeldt 2006 M119
Japanese 1.2% 2390 Sato 2014 M119

A 2008 study by Li suggested that the admixture analyses of Tai–Kadai-speaking populations showed a significant genetic influence in a large proportion of Indonesians. Most of the population samples contained a high frequency of haplogroup O-M119 (Hui 2008).

The frequencies of Haplogroup O-M119 among various East Asian and Austronesian populations suggest a complex genetic history of the modern Han populations of southern China.[citation needed] Although Haplogroup O-M119 occurs only at an average frequency of approximately 4% among Han populations of northern China and peoples of southwestern China and Southeast Asia who speak Tibeto-Burman languages, the frequency of this haplogroup among the Han populations of southern China nearly quadruples to about 15-23%.[4] The frequency of Haplogroup O-M119 among the Southern Han has been found to be slightly greater than the arithmetic mean of the frequencies of Haplogroup O-M119 among the Northern Han and a pooled sample of Austronesian populations. This suggests that modern Southern Han populations may possess a non-trivial number of male ancestors who were originally affiliated with some Austronesian-related culture, or who at least shared some genetic affinity with many of the ancestors of modern Austronesian peoples.[5][6]

Subclade distribution[edit]


This lineage is found frequently in Austronesians, southern Han Chinese, and Tai peoples.[7] This lineage is presumed to be a marker of the prehistoric Austronesian expansion, with possible origins encompassing the regions along the southeastern coast of China and neighboring Taiwan, and is found among modern populations of Maritime Southeast Asia and Oceania (Karafet 2005).

Haplogroup O-M119 Y-chromosomes also have been found to occur at low frequency among various populations of Siberia, such as the Nivkhs (one of 17 sampled Y-chromosomes), Ulchi/Nanai (2/53), Yenisey Evenks (1/31), and especially the Buryats living in the Sayan-Baikal uplands of Irkutsk Oblast (6/13) (Lell 2002).


O-P203 was found in 86.7% (52/60) of a sample from Nias, 70.8% (34/48) of Taiwanese Aboriginals, 28.4% (21/74) of Mentawai, 11.4% (73/641) of Balinese, 9.8% (6/61) of a sample from Java, 9.1% (36/394) of a sample from Flores, 9.1% (15/165) of Han Chinese, 8.3% (1/12) of a sample from Western Samoa, 8.2% (4/49) of Tujia from Hunan, 6.9% (4/58) of Miao from China, 5.7% (4/70) of Vietnamese, 3.3% (1/30) of a sample from the Moluccas, 3.1% (1/32) of Malaysians, 3.0% (1/33) of a sample from highland Papua New Guinea, 2.6% (1/38) of a sample from Sumatra, 2.3% (2/86) of a sample from Borneo, 2.1% (1/48) of Filipinos, 2.0% (1/51) of She, 1.7% (1/60) of Yao from Guangxi, 1.1% (1/92) of a sample from Lembata, and 0.9% (3/350) of a sample from Sumba.[8]

In a study published in 2011, O-P203 was observed in 22.2% (37/167) of Han Chinese male volunteers at Fudan University in Shanghai whose origin may be traced back to East China (Jiangsu, Zhejiang, Shanghai, or Anhui), 12.3% (8/65) of Han Chinese male volunteers whose origin may be traced back to South China, and 1.6% (2/129) of Han Chinese male volunteers whose origin may be traced back to North China.[9]


This lineage was observed in one individual from China (Underhill 2000) and another from Kota Kinabalu (Hurles 2005).


This lineage occurs among Austronesian peoples of Taiwan, the Philippines, Indonesia, Melanesia, Micronesia, and Madagascar as well as among some populations of continental Southeast Asia and among Bantu peoples of the Comoros.[10][citation needed] It also has been found in a Hawaiian.[11]

A study published in 2010 found O-M110 in 18.8% (9/48) Taiwanese Aboriginals, 13.3% (8/60) Nias, 8.3% (4/48) Philippines, 7.4% (4/54) Sulawesi, 6.3% (22/350) Sumba, 5.8% (5/86) Borneo, 3.3% (1/30) Moluccas, 2.3% (1/44) Maewo, Vanuatu, 1.6% (1/61) Java, 1.4% (1/74) Mentawai, and 0.8% (5/641) Bali.[12]

A study published in 2012 found O-M110 in 4.6% (33/712) of males from the Solomon Islands.[13]


Phylogenetic history[edit]

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
O-M175 26 VII 1U 28 Eu16 H9 I O* O O O O O O O O O O
O-M119 26 VII 1U 32 Eu16 H9 H O1* O1a O1a O1a O1a O1a O1a O1a O1a O1a O1a
O-M101 26 VII 1U 32 Eu16 H9 H O1a O1a1 O1a1a O1a1a O1a1 O1a1 O1a1a O1a1a O1a1a O1a1a O1a1a
O-M50 26 VII 1U 32 Eu16 H10 H O1b O1a2 O1a2 O1a2 O1a2 O1a2 O1a2 O1a2 O1a2 O1a2 O1a2
O-P31 26 VII 1U 33 Eu16 H5 I O2* O2 O2 O2 O2 O2 O2 O2 O2 O2 O2
O-M95 26 VII 1U 34 Eu16 H11 G O2a* O2a O2a O2a O2a O2a O2a O2a O2a O2a1 O2a1
O-M88 26 VII 1U 34 Eu16 H12 G O2a1 O2a1 O2a1 O2a1 O2a1 O2a1 O2a1 O2a1 O2a1 O2a1a O2a1a
O-SRY465 20 VII 1U 35 Eu16 H5 I O2b* O2b O2b O2b O2b O2b O2b O2b O2b O2b O2b
O-47z 5 VII 1U 26 Eu16 H5 I O2b1 O2b1a O2b1 O2b1 O2b1a O2b1a O2b1 O2b1 O2b1 O2b1 O2b1
O-M122 26 VII 1U 29 Eu16 H6 L O3* O3 O3 O3 O3 O3 O3 O3 O3 O3 O3
O-M121 26 VII 1U 29 Eu16 H6 L O3a O3a O3a1 O3a1 O3a1 O3a1 O3a1 O3a1 O3a1 O3a1a O3a1a
O-M164 26 VII 1U 29 Eu16 H6 L O3b O3b O3a2 O3a2 O3a2 O3a2 O3a2 O3a2 O3a2 O3a1b O3a1b
O-M159 13 VII 1U 31 Eu16 H6 L O3c O3c O3a3a O3a3a O3a3 O3a3 O3a3a O3a3a O3a3a O3a3a O3a3a
O-M7 26 VII 1U 29 Eu16 H7 L O3d* O3c O3a3b O3a3b O3a4 O3a4 O3a3b O3a3b O3a3b O3a2b O3a2b
O-M113 26 VII 1U 29 Eu16 H7 L O3d1 O3c1 O3a3b1 O3a3b1 - O3a4a O3a3b1 O3a3b1 O3a3b1 O3a2b1 O3a2b1
O-M134 26 VII 1U 30 Eu16 H8 L O3e* O3d O3a3c O3a3c O3a5 O3a5 O3a3c O3a3c O3a3c O3a2c1 O3a2c1
O-M117 26 VII 1U 30 Eu16 H8 L O3e1* O3d1 O3a3c1 O3a3c1 O3a5a O3a5a O3a3c1 O3a3c1 O3a3c1 O3a2c1a O3a2c1a
O-M162 26 VII 1U 30 Eu16 H8 L O3e1a O3d1a O3a3c1a O3a3c1a O3a5a1 O3a5a1 O3a3c1a O3a3c1a O3a3c1a O3a2c1a1 O3a2c1a1

Research publications[edit]

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

Phylogenetic trees[edit]

This phylogenetic tree of haplogroup O subclades is based on the YCC 2008 tree (Karafet 2008) and subsequent published research.

  • O-MSY2.2 (MSY2.2)
    • O-M119 (M119)
      • O-P203.2 (M307.2/P203.2)
      • O-M50 (M50, M103, M110)

See also[edit]


Y-DNA O subclades[edit]

Y-DNA backbone tree[edit]

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
F1  F2  F3  GHIJK
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
  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 4884158Freely 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.)



Works cited[edit]



Further reading[edit]


  1. ^ G. David Poznik, Yali Xue, Fernando L. Mendez, et al., "Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences." Nature Genetics 2016 June ; 48(6): 593–599. doi:10.1038/ng.3559.
  2. ^ a b YFull Haplogroup YTree v5.04 at 16 May 2017
  3. ^ https://isogg.org/tree/ISOGG_HapgrpO.html: "MSY2.2 was removed from tree because not providing reliable results."
  4. ^ Yan, Shi (September 2011). "An updated tree of Y-chromosome Haplogroup O and revised phylogenetic positions of mutations P164 and PK4". European Journal Of Genetics. 19: 1013–5. PMC 3179364Freely accessible. PMID 21505448. doi:10.1038/ejhg.2011.64. 
  5. ^ HUGO Pan-Asian SNP Consortium, HUGO Pan-Asian SNP Consortium. "Mapping Human Genetic Diversity in Asia". www.sciencemag.org. Science Magazine. Retrieved 11 December 2009. 
  6. ^ HUGO Pan-Asian SNP Consortium, HUGO Pan-Asian SNP Consortium. "Mapping Human Genetic Diversity in Asia". www.sciencemag.org. Science Magazine. Retrieved 11 December 2009. 
  7. ^ "Major East–West Division Underlies Y Chromosome Stratification across Indonesia". Karafet et al. Retrieved 2010.  Check date values in: |access-date= (help)
  8. ^ Karafet, Tatiana. "Major East–West Division Underlies Y Chromosome Stratification across Indonesia". http://mbe.oxfordjournals.org/. Oxford Journals. Retrieved 5 March 2010.  External link in |website= (help)
  9. ^ Shi Yan, Chuan-Chao Wang, Hui Li, Shi-Lin Li, Li Jin, and The Genographic Consortium, "An updated tree of Y-chromosome Haplogroup O and revised phylogenetic positions of mutations P164 and PK4." European Journal of Human Genetics (2011) 19, 1013–1015; doi:10.1038/ejhg.2011.64; published online 20 April 2011.
  10. ^ Said Msaidie, Axel Ducourneau, Gilles Boetsch, et al., "Genetic diversity on the Comoros Islands shows early seafaring as major determinant of human biocultural evolution in the Western Indian Ocean." European Journal of Human Genetics (2010), 1–6. doi:10.1038/ejhg.2010.128
  11. ^ Swapan Mallick, Heng Li, Mark Lipson, et al., "The Simons Genome Diversity Project: 300 genomes from 142 diverse populations." Nature 538, 201–206 (13 October 2016) doi:10.1038/nature18964
  12. ^ Karafet, T. M.; Hallmark, B.; Cox, M. P.; Sudoyo, H.; Downey, S.; Lansing, J. S.; Hammer, M. F. (2010). "Major East-West Division Underlies Y Chromosome Stratification across Indonesia". Molecular Biology and Evolution. 27 (8): 1833–44. PMID 20207712. doi:10.1093/molbev/msq063. 
  13. ^ Frederick Delfin, Sean Myles, Ying Choi, David Hughes, Robert Illek, Mannis van Oven, Brigitte Pakendorf, Manfred Kayser, and Mark Stoneking, "Bridging Near and Remote Oceania: mtDNA and NRY Variation in the Solomon Islands." Molecular Biology and Evolution 29(2):545–564. 2012 doi:10.1093/molbev/msr186.