Haplogroup O-M119

Haplogroup O-M119
Possible time of origin	33,103 [95% CI 24,460 <-> 40,854] years ago (Karmin 2015) 34,100 or 29,200 years ago (Poznik 2016) 30,100 [95% CI 27,800 <-> 32,400] years before present (YFull 2017)
Coalescence age	16,538 [95% CI 11,755 <-> 21,147] years ago (Karmin 2015) 15,000 [95% CI 13,200 <-> 16,900] years before present (YFull 2017)
Possible place of origin	Southeast Asia or Southern China
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.

Origins

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

Paleolithic migrations

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

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.

Distribution

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 MSY2.2(xM50)=1
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
Han (NE China)	4.8%	42	Katoh 2005	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)
Manchu	3.8%	52	Hammer 2006	M119
Koreans (Daejeon)	3.8%	133	Park 2012	P203=3 M119(xP203, M110)=2
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 (NE China)	3.0%	101	Katoh 2005	M119
Koreans (Seoul)	3.0%	573	Park 2012	P203=16 M119(xP203, M110)=1
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
Koreans	2.7%	300	Park 2013	M119
Koreans	2.7%	75	Hammer 2006	M119
Japanese (Kantō)	2.6%	117	Katoh 2005	M119
Koreans (Seoul)	2.4%	85	Katoh 2005	M119
Lavongai	2.3%	43	Scheinfeldt 2006	M119
Koreans (South Korea)	2.2%	506	Kim 2011	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
Mongolia (mostly Khalkh)	0.7%	149	Hammer 2006	M119
Mongols (Mongolia)	0.6%	160	DiCristofaro 2013	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. 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%. 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.

Subclade distribution

O-M119

This lineage is found frequently in Austronesians, southern Han Chinese, and Tai peoples. 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

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.

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.

O-M101

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

O-M50

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. It also has been found in a Hawaiian.

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.

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

Phylogenetics

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
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

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

3

Phylogenetic trees

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

• Austro-Tai languages

Genetics

3

Y-DNA O subclades

3

References

Works cited

Journals

• Chen, Chen; Hui, LI; Zhen-Dong, QIN; Wen-Hong, LIU; Wei-Xiong, LIN; Rui-Xing, YIN; Li, JIN; Shang-Ling, PAN (2006). "Y-chromosome Genotyping and Genetic Structure of Zhuang Populations". Acta Genetica Sinica. 33 (12): 1060–72. doi:10.1016/S0379-4172(06)60143-1. PMID 17185165.
• Hammer, Michael F.; Karafet, Tatiana M.; Park, Hwayong; Omoto, Keiichi; Harihara, Shinji; Stoneking, Mark; Horai, Satoshi (2005). "Dual origins of the Japanese: Common ground for hunter-gatherer and farmer Y chromosomes". Journal of Human Genetics. 51 (1): 47–58. doi:10.1007/s10038-005-0322-0. PMID 16328082.
• Hurles, M; Sykes, B; Jobling, M; Forster, P (2005). "The Dual Origin of the Malagasy in Island Southeast Asia and East Africa: Evidence from Maternal and Paternal Lineages". The American Journal of Human Genetics. 76 (5): 894–901. doi:10.1086/430051. PMC 1199379. PMID 15793703.
• Karafet, Tatiana M.; Lansing, J. S.; Redd, Alan J.; Watkins, Joseph C.; Surata, S. P. K.; Arthawiguna, W. A.; Mayer, Laura; Bamshad, Michael; et al. (2005). "Balinese Y-Chromosome Perspective on the Peopling of Indonesia: Genetic Contributions from Pre-Neolithic Hunter-Gatherers, Austronesian Farmers, and Indian Traders". Human Biology. 77 (1): 93–114. doi:10.1353/hub.2005.0030. PMID 16114819.
• 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. doi:10.1093/molbev/msq063. PMID 20207712.
• Kayser, Manfred; Brauer, Silke; Weiss, Gunter; Schiefenhövel, Wulf; Underhill, Peter; Shen, Peidong; Oefner, Peter; Tommaseo-Ponzetta, Mila; Stoneking, Mark (2003). "Reduced Y-Chromosome, but Not Mitochondrial DNA, Diversity in Human Populations from West New Guinea". The American Journal of Human Genetics. 72 (2): 281–302. doi:10.1086/346065. PMC 379223. PMID 12532283.
• Kayser, M.; Choi, Y.; Van Oven, M.; Mona, S.; Brauer, S.; Trent, R. J.; Suarkia, D.; Schiefenhovel, W.; Stoneking, M. (2008). "The Impact of the Austronesian Expansion: Evidence from mtDNA and Y Chromosome Diversity in the Admiralty Islands of Melanesia". Molecular Biology and Evolution. 25 (7): 1362–74. doi:10.1093/molbev/msn078. PMID 18390477.
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Websites

• ISOGG (2010). "Y-DNA Haplogroup O and its Subclades. 2010". International Society of Genetic Genealogy. Retrieved 2010. {{cite web}}: Check date values in: |accessdate= (help)