Haplogroup O-M119

Haplogroup O-M119
Possible time of origin 33,103 [95% CI 24,460 <-> 40,854] years ago (Karmin 2015[1])

34,100 or 29,200 years ago (Poznik 2016[2])

30,100 [95% CI 27,800 <-> 32,400] years before present (YFull 2017[3])
Possible place of origin Southeast Asia or Southern China
Coalescence age 16,538 [95% CI 11,755 <-> 21,147] years ago (Karmin 2015[1])

15,000 [95% CI 13,200 <-> 16,900] years before present (YFull 2017[3])
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.[4]

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
Nias100.0%60Karafet 2010M119
Nias99.8%407vanOven 2011M119
Taiwanese aborigines89.6%48Karafet 2010M119
Mentawai86.5%74Karafet 2010M119
Aboriginal Taiwanese83.4%223Tajima 2004M119
Taiwan (aborigines)71.8%39Hurles 2005M119(xM101)
Taiwan (aborigines)68.9%74Underhill 2000M119(xM101)
Hlai/Cun58%-Li 2008 -
Tagalog
(Philippine subgroup)		46.0%50Tajima 2004M119
Philippines35.7%28Hurles 2005M119(xM101)
Kota Kinabalu29.2%65Hurles 2005M119
Trobriand Islands28%-Kayser 2003 -
Li (Hlai)26.5%34Xue 2006M119
Malay (near Kuala Lumpur)25.0%12Tajima 2004M119
Han (East China)24.0%167Yan 2011M119
Han Chinese23%-Kayser 2003 -
Java23%-Kayser 2003 -
Nusa Tenggara23%-Kayser 2003 -
Banjarmasin22.7%22Hurles 2005M119(xM101)
Balinese21.1%641Karafet 2010M119
Mandar20.4%54Karafet 2010M119(xP203)
Tujia20%-Su 1999-
Han (Meixian)20.0%35Xue 2006M119
Buka20.0%10Scheinfeldt 2006M119
Thin Board Mien18.2%11Cai 2011M119(xM110)
Majuro (Marshall Islands)18.2%11Hurles 2005M50
Admiralty Islands18%-Kayser 2008 -
Balinese18%-Karafet 2005 -
Sui18%-Xie 2004 -
Zhuang18%-Su 1999 -
Malagasy17.1%35Hurles 2005M50
Han (South China)16.9%65Yan 2011M119
Qiang15.2%33Xue 2006M119
Borneo15%-Kayser 2003-
Han Chinese15%-Tajima 2004-
Java14.8%61Karafet 2010M119
She14.7%34Xue 2006M119
Han (Chengdu)14.7%34Xue 2006M119
Manus14.3%7Scheinfeldt 2006M119
Palyu13.3%30Cai 2011M119
Batak Toba13.2%38Karafet 2010M119(xM110)
Sumba12.6%350Karafet 2010M119
Micronesia12.5%16Karafet 2010M119(xP203, M110)
Guizhou Miao12.2%49Cai 2011M119(xM110)
Lowland Kimmun12.2%41Cai 2011M119(xM110)
Thai (Northern Thailand)11.8%17He 2012P203(xM101)
Bunu11.1%36Cai 2011M119(xM110)
Filipinos10.4%48Karafet 2010M119
Zhuang10%-Hammer 2006-
Mountain Straggler Mien10.0%20Cai 2011M119(xM110)
Han (China & Taiwan)9.7%165Karafet 2010M119(xM110)
Flores9.6%394Karafet 2010M119(xM110)
Zhuang9.6%166Chen 2006-
Han (Taiwan)9.5%21Tajima 2004M119
Han (Yili)9.4%32Xue 2006M119
Borneo (Indonesia)9.3%86Karafet 2010M119
Bougainville9.2%65Scheinfeldt 2006M119
Top Board Mien9.1%11Cai 2011M119(xM110)
Northern Mien9.1%33Cai 2011M119(xM110)
Northern She8.9%56Cai 2011M119(xM110)
Thai
(Chiang Mai & Khon Kaen)		8.8%34Tajima 2004M119
Hui8.6%35Xue 2006M119
Mosuo (Ninglang, Yunnan)8.5%47Wen 2004M119(xM110)
Tonga8.3%12Karafet 2010M119(xP203, M110)
Tujia (Jishou, Hunan)8.2%49Karafet 2010P203
Hlai/Cun8%-Li 2008-
Ewenki (China)7.7%26Xue 2006M119
Xibe7.3%41Xue 2006M119
Hunan Miao7.0%100Cai 2011M119(xM110)
Tujia7%-Xie 2004-
Miao (China)6.9%58Karafet 2010P203
Katu6.7%45Cai 2011M119(xM110)
Moluccas6.7%30Karafet 2010M119
Han (Lanzhou)6.7%30Xue 2006M119
Kinh6.7%15Cai 2011M119(xM110)
Kinh (Hanoi, Vietnam)6.6%76He 2012P203(xM101)
Southern Mien6.5%31Cai 2011M119(xM110)
Malaysia6.3%32Karafet 2010M119(xM110)
Yunnan Miao6.1%49Cai 2011M119(xM110)
Northern Han6.1%49Tajima 2004M119
Comorians6.0%381Msaidie 2010M50=22
MSY2.2(xM50)=1
Bai (Dali, Yunnan)6.0%50Wen 2004M119(xM110)
Kyrgyz (Kyrgyzstan)5.8%52Wells 2001M119
Vietnam5.7%70Karafet 2010P203
Yao (Liannan, Guangdong)5.7%35Xue 2006M119
Samoa5.6%18Karafet 2010P203
Daur5.1%39Xue 2006M119
Cham
(Binh Thuan, Vietnam)		5.1%59He 2012M119(xM50)
Dungan (Kyrgyzstan)5.0%40Wells 2001M119
Han (NE China)4.8%42Katoh 2005M119
Maewo (Vanuatu)4.5%44Karafet 2010M119(xP203)
Korean4.4%45Wells 2001M119
Western Mien4.3%47Cai 2011M119(xM110)
Pumi (Ninglang, Yunnan)4.3%47Wen 2004M119(xM110)
Mongolian4.2%24Wells 2001M119
Western Samoa4.0%25Hurles 2005M119(xM101, M50)
Manchu3.8%52Hammer 2006M119
Koreans (Daejeon)3.8%133Park 2012P203=3
M119(xP203, M110)=2
New Ireland3.7%109Scheinfeldt 2006M119
Bo3.6%28Cai 2011M119(xM110)
Japanese3.4%263Nonaka 2007M119(xM101, M50)
Lembata3.3%92Karafet 2010M119(xM110)
Korean3.2%216Kim 2007M119
Han (North China)3.1%129Yan 2011M119(xM110)
Papua New Guinea
(Highlands)		3.0%33Karafet 2010P203
Manchu (NE China)3.0%101Katoh 2005M119
Koreans (Seoul)3.0%573Park 2012P203=16
M119(xP203, M110)=1
Manchu2.9%35Xue 2006M119
Han (Harbin)2.9%35Xue 2006M119
Buyi2.9%35Xue 2006M119
Yao (Bama, Guangxi)2.9%35Xue 2006M119
West New Britain2.8%249Scheinfeldt 2006M119
Koreans2.7%300Park 2013M119
Koreans2.7%75Hammer 2006M119
Japanese (Kantō)2.6%117Katoh 2005M119
Koreans (Seoul)2.4%85Katoh 2005M119
Lavongai2.3%43Scheinfeldt 2006M119
Koreans (South Korea)2.2%506Kim 2011M119
Laven2.0%50Cai 2011M119(xM110)
Yi (Shuangbai, Yunnan)2.0%50Wen 2004M119(xM110)
Hmong Daw (Laos)2.0%51Cai 2011M119(xM110)
She2.0%51Karafet 2010P203
Japanese (Kyushu)1.9%104Tajima 2004M119
Vanuatu1.9%52Hurles 2005M50
Yao (Guangxi)1.7%60Karafet 2010P203
Uygur1.4%70Xue 2006M119
East New Britain1.4%145Scheinfeldt 2006M119
Japanese1.2%2390Sato 2014M119
Mongolia
(mostly Khalkh)		0.7%149Hammer 2006M119
Mongols (Mongolia)0.6%160DiCristofaro 2013M119

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%.[5] 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.[6][7]

Subclade distribution

O-M119

This lineage is found frequently in Austronesians, southern Han Chinese, and Tai peoples.[8] 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.[9]

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.[10]

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.[11] It also has been found in a Hawaiian.[12]

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.[13]

A study published in 2012 found O-M110 in 4.6% (33/712) of males from the Solomon Islands.[14]

Phylogenetics

Phylogenetic history

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-M17526VII1U28Eu16H9IO*OOOOOOOOOO
O-M11926VII1U32Eu16H9HO1*O1aO1aO1aO1aO1aO1aO1aO1aO1aO1a
O-M10126VII1U32Eu16H9HO1aO1a1O1a1aO1a1aO1a1O1a1O1a1aO1a1aO1a1aO1a1aO1a1a
O-M5026VII1U32Eu16H10HO1bO1a2O1a2O1a2O1a2O1a2O1a2O1a2O1a2O1a2O1a2
O-P3126VII1U33Eu16H5IO2*O2O2O2O2O2O2O2O2O2O2
O-M9526VII1U34Eu16H11GO2a*O2aO2aO2aO2aO2aO2aO2aO2aO2a1O2a1
O-M8826VII1U34Eu16H12GO2a1O2a1O2a1O2a1O2a1O2a1O2a1O2a1O2a1O2a1aO2a1a
O-SRY46520VII1U35Eu16H5IO2b*O2bO2bO2bO2bO2bO2bO2bO2bO2bO2b
O-47z5VII1U26Eu16H5IO2b1O2b1aO2b1O2b1O2b1aO2b1aO2b1O2b1O2b1O2b1O2b1
O-M12226VII1U29Eu16H6LO3*O3O3O3O3O3O3O3O3O3O3
O-M12126VII1U29Eu16H6LO3aO3aO3a1O3a1O3a1O3a1O3a1O3a1O3a1O3a1aO3a1a
O-M16426VII1U29Eu16H6LO3bO3bO3a2O3a2O3a2O3a2O3a2O3a2O3a2O3a1bO3a1b
O-M15913VII1U31Eu16H6LO3cO3cO3a3aO3a3aO3a3O3a3O3a3aO3a3aO3a3aO3a3aO3a3a
O-M726VII1U29Eu16H7LO3d*O3cO3a3bO3a3bO3a4O3a4O3a3bO3a3bO3a3bO3a2bO3a2b
O-M11326VII1U29Eu16H7LO3d1O3c1O3a3b1O3a3b1-O3a4aO3a3b1O3a3b1O3a3b1O3a2b1O3a2b1
O-M13426VII1U30Eu16H8LO3e*O3dO3a3cO3a3cO3a5O3a5O3a3cO3a3cO3a3cO3a2c1O3a2c1
O-M11726VII1U30Eu16H8LO3e1*O3d1O3a3c1O3a3c1O3a5aO3a5aO3a3c1O3a3c1O3a3c1O3a2c1aO3a2c1a
O-M16226VII1U30Eu16H8LO3e1aO3d1aO3a3c1aO3a3c1aO3a5a1O3a5a1O3a3c1aO3a3c1aO3a3c1aO3a2c1a1O3a2c1a1

Research publications

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

Phylogenetic trees

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

See also

Genetics

Y-DNA O subclades

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
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

Footnotes

    Works cited

    Journals

    Websites

    References

    1. 1 2 Monika Karmin, Lauri Saag, Mário Vicente, et al., "A recent bottleneck of Y chromosome diversity coincides with a global change in culture." Genome Research (2015) 25: 459-466. doi: 10.1101/gr.186684.114
    2. ↑ 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.
    3. 1 2 YFull Haplogroup YTree v5.04 at 16 May 2017
    4. ↑ https://isogg.org/tree/ISOGG_HapgrpO.html: "MSY2.2 was removed from tree because not providing reliable results."
    5. ↑ 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 3179364 Freely accessible. PMID 21505448. doi:10.1038/ejhg.2011.64.
    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. ↑ 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.
    8. ↑ "Major East–West Division Underlies Y Chromosome Stratification across Indonesia". Karafet et al. Retrieved 2010. Check date values in: |access-date= (help)
    9. ↑ 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)
    10. ↑ 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.
    11. ↑ 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
    12. ↑ 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
    13. ↑ 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.
    14. ↑ 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.
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