Haplogroup B (mtDNA)

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Haplogroup B
Peopling of eurasia.jpg
Possible time of origin50,000 YBP
Possible place of originSouth Asia
AncestorR11'B
DescendantsB4, B5, B7
Defining mutations8281-8289d[1]

In human mitochondrial genetics, haplogroup B is a human mitochondrial DNA (mtDNA) haplogroup.

Origin[edit]

Haplogroup B is believed to have arisen in Asia some 50,000 years before present. Its ancestral haplogroup was haplogroup R.

The greatest variety of haplogroup B is in China. It is therefore likely that it underwent its earliest diversification in mainland East or South East Asia.[2]

Distribution[edit]

Basal B was found in Upper Paleolithic Tianyuan man.[3]

Haplogroup B is now most common among populations native to Southeast Asia,[4] as well as speakers of Sino-Tibetan languages and Austronesian languages.

A subclade of B4b (which is sometimes labeled B2) is one of five haplogroups found among the indigenous peoples of the Americas, the others being A, C, D, and X.

Because the migration to the Americas by the ancestors of indigenous Americans is generally believed to have been from northeastern Siberia via Beringia, it is surprising that Haplogroup B and Haplogroup X have not been found in Paleo-Siberian tribes of northeastern Siberia.[5] However, Haplogroup B has been found among Turkic, Mongolic, and Tungusic populations of Siberia, such as Tuvans, Altays, Shors, Khakassians, Yakuts, Buryats, Mongols, Negidals, and Evenks.[6] This haplogroup is also found among populations in China, Indonesia, Iran,[6] Iraq,[7] Japan, Korea, Laos,[8] Madagascar, Malaysia, Melanesia, Micronesia, Mongolia, the Philippines, Polynesia, Thailand, and Vietnam.[9][10]

Although haplogroup B in general has been found in many Siberian population samples, the subclade that is phylogenetically closest to American B2, namely B4b1, has been found mainly in populations of southern China and Southeast Asia, especially Filipinos and Austronesian speakers of eastern Indonesia (approx. 8%) and the aborigines of Taiwan and Hainan (approx. 7%).[11][12][13] However, B4b1 has been observed in populations as far north as Turochak and Choya districts in the north of Altai Republic (3/72 = 4.2% Tubalar),[14] Miyazaki and Tokyo, Japan (approx. 3%),[15] South Korea (4/185 = 2.2%),[10] Tuva (1/95 = 1.1% Tuvan),[14] and Hulunbuir (1/149 = 0.7% Barghut).[16]

Table of Frequencies of MtDNA Haplogroup B[edit]

Population Frequency Count Source Subtypes
Carolinian (Saipan) 1.000 17 [citation needed] B4a1a1a=15, B4a1a(xB4a1a1a)=2
Dingban Yao (Mengla, Yunnan) 0.600 10 [citation needed] B4(xB4a, B4b1, B4e, B4f)=3, B4a=1, B4e=1, B4f=1
Iu Mien (Mengla, Yunnan) 0.481 27 [citation needed] B4a=7, B4b1=5, B5a=1
Bapai Yao (Liannan, Guangdong) 0.429 35 [citation needed] B5a=7, B4e=3, B4b1=2, B4a=1, B4(xB4a, B4b1, B4e, B4f)=1, B(xB4, B5)=1
Filipino 0.422 64 [11] B4a1a(xB4a1a1)=8, B5b=7, B4b1=6, B4c1b=4, B4a1a1=1, B5a=1
Tu Yao (Hezhou, Guangxi) 0.390 41 [citation needed] B5a=7, B4(xB4a, B4b1, B4e, B4f)=6, B4a=3
Guoshan Yao (Jianghua, Hunan) 0.375 24 [citation needed] B5a=7, B4(xB4a, B4b1, B4e, B4f)=2
Cham (Bình Thuận, Vietnam) 0.369 168 [17] B5a=27, B4c2=18, B4c1b2=3, B4h=3, B4a1(xB4a1a)=2, B4g=2, B5b2a=2, B4a(xB4a1)=1, B4a1a=1, B4b1=1, B4c1b(xB4c1b2)=1, B5b1=1
Huatou Yao (Fangcheng, Guangxi) 0.368 19 [citation needed] B5a=3, B4a=2, B4b1=1, B4e=1
Filipino (Luzon) 0.367 177 [11] B4a1a(xB4a1a1)=20, B5b=18, B4b1=13, B4c1b=10, B5a=2, B4a(xB4a1a)=1, B7=1
Hmong (Wenshan, Yunnan) 0.333 39 [citation needed] B5a=8, B4a=5
Dong (Tianzhu County, Guizhou) 0.321 28 [citation needed] B4a=6, B5a=2, B4c=1
Kinh (Vietnamese) 0.317 139 [citation needed] B4=24, B5=19, B6=1
Mien (Shangsi, Guangxi) 0.313 32 [citation needed] B4a=5, B5a=3, B4e=2
Kim Mun (Malipo, Yunnan) 0.300 40 [citation needed] B5a=5, B4a=3, B4e=2, B4b1=1, B5(xB5a, B5b)=1
Dai (Xishuangbanna, Yunnan) 0.293 41 [citation needed] B5a=4, B(xB4, B5a, B5b)=4, B4a=3, B4(xB4a)=1
Bunu (Dahua & Tianlin, Guangxi) 0.280 25 [citation needed] B4a=5, B5a=2
Filipino (Visayas) 0.277 112 [11] B4a1a(xB4a1a1)=11, B4b1=8, B4c1b=7, B5b=5
Lanten Yao (Tianlin, Guangxi) 0.269 26 [citation needed] B4a=3, B5a=2, B4b1=1, B4f=1
Laos 0.262 214 [8] B5a=26, B4g=7, B4c2=6, B4a'g*=3, B4a1*=3, B4b1a2a=2, B4e=2, B4*=1, B4a1b=1, B4b1*=1, B4b1a1'2*=1, B4c1b=1, B5b=1, B6=1
Wuzhou Yao (Fuchuan, Guangxi) 0.258 31 [citation needed] B4a=4, B5a=3, B4b1=1
Lahu (Simao, Yunnan) 0.250 32 [citation needed] B4a=5, B4(xB4a, B4b1)=3
Pan Yao (Tianlin, Guangxi) 0.250 32 [citation needed] B5a=4, B4a=2, B4f=1, B5b=1
Yi (Xishuangbanna, Yunnan) 0.250 16 [citation needed] B4(xB4a, B4b1)=1, B4a=1, B5(xB5a, B5b)=1, B5a=1
Naxi (Lijiang, Yunnan) 0.244 45 [citation needed] B4a=5, B4(xB4a, B4b1)=3, B5a=3
Filipino (Mindanao) 0.243 70 [11] B4a1a(xB4a1a1)=7, B4b1=5, B5b=3, B4a(xB4a1a)=1, B4a1a1=1
Lowland Yao (Fuchuan, Guangxi) 0.238 42 [citation needed] B4a=3, B4b1=3, B4(xB4a, B4b1, B4e, B4f)=2, B5a=1, B5b=1
Han (Taiwan) 0.234 111 [citation needed] B4+C16261T+G16129A!=6, B5a1=3, B4+C16261T=2, B4b1a2=2, B4b1b'c=2, B4c1b2a(xB4c1b2a2)=2, B4g=2, B4a1a(xB4a1a1a)=1, B4a4=1, B4b1a3=1, B4c1b+A16335G=1, B4c1c=1, B5b(xB5b2a)=1, B5b2a=1
Gelao (Daozhen County, Guizhou) 0.226 31 [citation needed] B4a=3, B4(xB4a, B4b, B4c)=2, B4b=1, B5a=1
Bai (Xishuangbanna, Yunnan) 0.211 19 [citation needed] B4(xB4a, B4b1)=2, B4a=1, B5a=1
Han (Taipei, Taiwan) 0.209 91 [citation needed] B4(xB4f)=13, B5=6
Gelao (Daozhen County, Guizhou) 0.206 102 [citation needed] B4a=5, B4b1=5, B5a=5, B4(xB4a, B4b, B4c, B4e, B4f, B4g)=3, B5b=2, B4c=1
Korean (South Korea) 0.204 103 [6] B4=13, B5=8
CHS (Han from Hunan & Fujian) 0.200 55 [citation needed] B5=6, B4=5
Lahu (Xishuangbanna, Yunnan) 0.200 15 [citation needed] B4(xB4a, B4b1)=1, B4a=1, B4b1=1
Han (Southern California) 0.192 390 [citation needed] B=75
Dai (Xishuangbanna, Yunnan) 0.190 21 [citation needed] B4a=2, B4(xB4a)=1, B5b=1
Han (Southwest China; pool of 44 Sichuan, 34 Chongqing, 33 Yunnan, and 26 Guizhou) 0.190 137 [citation needed] B4b'd=9, B4a=6, B5a=4, B4c=3, B(xB4a, B4b'd, B4c, B5a, B5b, B6)=2, B5b=1, B6=1
Tujia (western Hunan) 0.188 64 [citation needed] B4a=5, B5b=3, B4(xB4a, B4b1)=2, B5a=2
Hmong (Jishou, Hunan) 0.184 103 [citation needed] B5a=6, B4a=5, B4b1=3, B4(xB4a, B4b1, B4e, B4f)=2, B4e=1, B4f=1, B5b=1
Hani (Xishuangbanna, Yunnan) 0.182 33 [citation needed] B5a=2, B4(xB4a, B4b1)=1, B4a=1, B4b1=1, B5b=1
Xiban Yao (Fangcheng, Guangxi) 0.182 11 [citation needed] B4a=1, B4b1=1
Korean (Seoul & Daejeon, South Korea) 0.180 261 Kim 2008 B(xB5)=32, B5=15
Han (Taiwan) 0.179 1117 [citation needed] B=200
CHD (Han from Denver, Colorado) 0.178 73 [citation needed] B4=12, B5=1
Hui (Xinjiang) 0.178 45 [citation needed] B4a=4, B5a=2, B4b1=1, B6=1
Yi (Shuangbai, Yunnan) 0.175 40 [citation needed] B4(xB4a, B4b1)=5, B4a=1, B4b1=1
Tujia (Yanhe County, Guizhou) 0.172 29 [citation needed] B4b=3, B5a=2
Thailand 0.171 105 [citation needed] B(xB5a1)=13, B5a1=5
Tibetan (Zhongdian, Yunnan) 0.171 35 [citation needed] B5a=4, B4(xB4a, B4b1)=2
Aini (Xishuangbanna, Yunnan) 0.160 50 [citation needed] B4a=4, B(xB4, B5)=2, B4(xB4a, B4b1)=1, B5a=1
Chinese (Shenyang, Liaoning) 0.156 160 [citation needed] B4(xB4f)=15, B5=10
Daur (Evenk Autonomous Banner) 0.156 45 [citation needed] B4(xB4a, B4b)=4, B4a=3
Korean (Seoul National University Hospital) 0.155 633 [citation needed] B=98
Yi (Hezhang County, Guizhou) 0.150 20 [citation needed] B4b=1, B4(xB4a, B4b, B4c)=1, B6=1
Mongolian (Ulan Bator) 0.149 47 [6] B4=5, B5=2
Korean (South Korea) 0.148 203 [citation needed] B4(xB4f)=24, B5=6
Korean (South Korea) 0.146 185 [10] B4a=11, B4(xB4a, B4b, B4c)=7, B4b1=4, B5a=2, B5b=2, B4c=1
Vietnamese 0.143 42 [10] B4(xB4a, B4b, B4c)=3, B4a=1, B4b(xB4b1)=1, B5a=1
Okinawa 0.141 326 [citation needed] B4(xB4f)=28, B4f=10, B5=8
CHB (Han from Beijing Normal University) 0.140 121 [citation needed] B4=14, B5=3
JPT (Japanese from Tokyo) 0.136 118 [citation needed] B4=11, B5=5
Tujia (Yongshun, Hunan) 0.133 30 [citation needed] B4(xB4a, B4b1)=2, B5a=2
Japanese (Hokkaidō) 0.129 217 [citation needed] B4(xB4f)=18, B4f=5, B5=5
Japanese (Tōkai) 0.124 282 [citation needed] B4(xB4f)=26, B5=9
Japanese (Gifu) 0.121 1617 [citation needed] B=196
Negidal 0.121 33 [14] B5b2=4
Japanese (Tōhoku) 0.119 336 [citation needed] B4(xB4f)=24, B5=15, B4f=1
Korean (northern China) 0.118 51 [10] B4a=2, B4b(xB4b1)=2, B5a=1, B5b=1
Telengit (Altai Republic) 0.113 71 [6] B4=8
Jino (Xishuangbanna, Yunnan) 0.111 18 [citation needed] B4a=1, B5a=1
Tibetan (Nyingchi, Tibet) 0.111 54 [citation needed] B4a=5, B5b=1
Japanese (northern Kyūshū) 0.109 256 [citation needed] B4(xB4f)=19, B5=8, B4f=1
Japanese 0.109 211 [citation needed] B4b1=14, B5b=4, B5a=2, B4a=1, B4(xB4a, B4b1)=1, B5(xB5a, B5b)=1
Evenk (New Barag Left Banner) 0.106 47 [citation needed] B4b=4, B5b=1
Korean (Arun Banner) 0.104 48 [citation needed] B4b=2, B5b=2, B4(xB4a, B4b)=1
Mongol (New Barag Left Banner) 0.104 48 [citation needed] B4b=3, B4a=1, B5a=1
Han (Beijing) 0.100 40 [10] B(xB4, B5a, B5b)=2, B4b(xB4b1)=2
Nu (Gongshan, Yunnan) 0.100 30 [citation needed] B4(xB4a, B4b1)=3
Thai 0.100 40 [10] B5a=3, B(xB4, B5a, B5b)=1
Yi (Luxi, Yunnan) 0.097 31 [citation needed] B4(xB4a, B4b1)=2, B(xB4, B5)=1
Japanese (Miyazaki) 0.090 100 [15] B4b1a1=3, B4(xB4a, B4b1)=2, B4a=2, B5b=2
Khakassian (Khakassia) 0.088 57 [6] B4=5
Han (Xinjiang) 0.085 47 [citation needed] B5a=2, B4b1=1, B5b=1
Kyrgyz (Sary-Tash, Kyrgyzstan) 0.085 47 [citation needed] B5a=2, B4(xB4a, B4b1, B4c)=1, B4a=1
Mongolian (Ulan Bator) 0.085 47 [10] B4(xB4a, B4b, B4c)=2, B4a=1, B5b=1
Tuvan 0.084 95 [14] B4a1c2=4, B5=2, B4(xB4a1c2, B4b1a, B4d1a)=1, B4b1a=1
Tibetan (Nyingchi, Tibet) 0.083 24 [citation needed] B=2
Barghut (Hulunbuir) 0.081 149 [16] B4c1a2(xB4c1a2a)=4, B4f1=2, B5b(xB5b2)=2, B4c1a2a=1, B4b1a3a1a=1, B4d1=1, B5b2=1
Chamorro (85 Guam, 14 Saipan, & 6 Rota) 0.076 105 [citation needed] B4a1a1a=7, B4b1=1
Manchurian 0.075 40 [10] B4(xB4a, B4b, B4c)=2, B4a=1
Uzbek (Uzbekistan/Kyrgyzstan) 0.075 40 [citation needed] B=3
Uyghur (Penjim, Panfilov District, Almaty Province, Kazakhstan) 0.073 55 [citation needed] B5a=3, B4(xB4a, B4b1, B4c)=1
Buryat 0.071 126 [citation needed] B4(xB4a, B4b)=5, B5b=2, B(xB4, B5a, B5b)=1, B4b=1
Khamnigan (Buryatia) 0.071 99 [6] B4=5, B5=2
Dungan (Uzbekistan/Kyrgyzstan) 0.063 16 [citation needed] B=1
Persian (eastern Iran) 0.061 82 [6] B4=4, B5=1
Nogai (Dagestan) 0.061 33 [citation needed] B=2
Tibetan (Nagchu, Tibet) 0.057 35 [citation needed] B=2
Kazakh (Kegen Valley, Almaty Province, Kazakhstan) 0.055 55 [citation needed] B4(xB4a, B4b1, B4c)=1, B4a=1, B5b=1
Lisu (Gongshan, Yunnan) 0.054 37 [citation needed] B4(xB4a, B4b1)=1, B5(xB5a, B5b)=1
Tharu (Chitwan, Nepal) 0.053 133 [citation needed] B5a=7
Uzbek (Xinjiang) 0.052 58 [citation needed] B4b1=2, B4c=1
Bukharan Arab (Uzbekistan/Kyrgyzstan) 0.050 20 [citation needed] B=1
Filipino (Palawan) 0.050 20 [citation needed] B4a1a=1
Tajik (Uzbekistan/Kyrgyzstan) 0.050 20 [citation needed] B=1
Turkmen (Uzbekistan/Kyrgyzstan) 0.050 20 [citation needed] B=1
Shor (Kemerovo Oblast) 0.049 82 [6] B4=4
Va (Simao, Yunnan) 0.045 22 [citation needed] B4(xB4a)=1
Altai Kizhi 0.044 90 [6] B4=3, B5=1
Bai (Dali, Yunnan) 0.044 68 [citation needed] B5a=2, B4b1=1
Tofalar 0.043 46 [14] B4a1c2=2
Kyrgyz (Bakay-Ata, Kyrgyzstan) 0.042 48 [citation needed] B4(xB4a, B4b1, B4c)=1, B5b=1
Tubalar 0.042 72 [14] B4b1a=3
Evenk (Krasnoyarsk Krai) 0.041 73 [6] B4=3
Kazakh (Kosh-Agach, Altai Republic) 0.041 98 [16] B5b2=3, B4c1b=1
Buryat (Kushun, Nizhneudinsky, Irkutsk Oblast) 0.040 25 [14] B4d1a=1
Kazakh (Xinjiang) 0.038 53 [citation needed] B4a=2
Teleut (Kemerovo Oblast) 0.038 53 [6] B4=2
Kalmyk (Kalmykia) 0.036 110 [6] B4=3, B5=1
Tibetan (Chamdo, Tibet) 0.034 29 [citation needed] B4a=1
Buryat (Buryatia) 0.034 295 [6] B4=9, B5=1
Lahu (Lancang, Yunnan) 0.029 35 [citation needed] B4(xB4a, B4b1)=1
Pumi (Ninglang, Yunnan) 0.028 36 [citation needed] B4(xB4a, B4b1)=1
Va (Ximeng & Gengma, Yunnan) 0.028 36 [citation needed] B5a=1
Yakut (Yakutia) 0.028 36 [6] B4=1
Tharu (Morang, Nepal) 0.025 40 [citation needed] B5a=1
Tibetan (Deqin, Yunnan) 0.025 40 [citation needed] B4(xB4a, B4b1)=1
Oroqen (Oroqen Autonomous Banner) 0.023 44 [citation needed] B4(xB4a, B4b)=1
Uyghur (Xinjiang) 0.021 47 [citation needed] B(xB4, B5, B6)=1
Oirat Mongol (Xinjiang) 0.020 49 [citation needed] B4b1=1
Ainu 0.020 51 [citation needed] B4f1=1
Tuvinian (Tuva) 0.019 105 [6] B4=2
Tibetan (Qinghai) 0.018 56 [citation needed] B4a=1
Nogai (Nogaysky, Dagestan & Adyge-Khabalsky, Karachay-Cherkessia) 0.015 206 [citation needed] B=3
Tibetan (Shannan, Tibet) 0.014 74 [citation needed] B4a=1
Yakut 0.012 423 [citation needed] B4(xB4b1)=3
B4b1=1
B5b2=1
Bashkir 0.009 221 [citation needed] B=2
Chukchi (Anadyr) 0.000 15 [6]
Uyghur (Uzbekistan/Kyrgyzstan) 0.000 16 [citation needed]
Crimean Tatar (Uzbekistan/Kyrgyzstan) 0.000 20 [citation needed]
Iranian (Uzbekistan/Kyrgyzstan) 0.000 20 [citation needed]
Karakalpak (Uzbekistan/Kyrgyzstan) 0.000 20 [citation needed]
Kazakh (Uzbekistan/Kyrgyzstan) 0.000 20 [citation needed]
Kyrgyz (Uzbekistan/Kyrgyzstan) 0.000 20 [citation needed]
Hindu (Chitwan, Nepal) 0.000 24 [citation needed]
Nganasan 0.000 24 [14]
Tibetan (Diqing, Yunnan) 0.000 24 [citation needed]
Kurd (northwestern Iran) 0.000 25 [6]
Andhra Pradesh (tribal) 0.000 29 [citation needed]
Tibetan (Shigatse, Tibet) 0.000 29 [citation needed]
Batak (Palawan) 0.000 31 [citation needed]
Ket 0.000 38 [14]
Tajik (Tajikistan) 0.000 44 [6]
Tibetan (Lhasa, Tibet) 0.000 44 [citation needed]
Evenk (Buryatia) 0.000 45 [6]
Udege (Gvasiugi, Imeni Lazo, Khabarovsk Krai) 0.000 46 [14]
Itelmen 0.000 47 [14]
Chuvash 0.000 55 [citation needed]
Nivkh (northern Sakhalin) 0.000 56 [14]
Komi-Zyryans 0.000 62 [citation needed]
Mansi 0.000 63 [citation needed]
Chukchi 0.000 66 [14]
Evenk (53 Stony Tunguska basin & 18 Tuguro-Chumikan) 0.000 71 [14]
Tatar (Aznakayevo) 0.000 71 [citation needed]
Komi-Permyaks 0.000 74 [citation needed]
Siberian Eskimo 0.000 79 [14]
Ulchi 0.000 87 [14]
Mansi 0.000 98 [14]
Udmurt 0.000 101 [citation needed]
Mordvinian 0.000 102 [citation needed]
Khanty 0.000 106 [citation needed]
Yakut 0.000 117 [citation needed]
Tatar (Buinsk) 0.000 126 [citation needed]
Mari 0.000 136 [citation needed]
Koryak 0.000 155 [14]
Tatar 0.000 228 [citation needed]

Subclades[edit]

Tree[edit]

This phylogenetic tree of haplogroup B subclades is based on the paper by Mannis van Oven and Manfred Kayser Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation[1] and subsequent published research.

  • B'R11'R24
    • B4'5 – China (Han from Zhanjiang, Paleolithic remains from Tianyuan Cave), the Philippines (Ivatan)
      • B4
        • B4-T16217C* – Vietnam (Lô Lô), Japan
        • B4a'g'h'i'k'm (B4-C16261T)
          • B4-C16261T* – Thailand (Lao Isan in Chaiyaphum Province[18]), Cambodia (Takeo), Vietnam (Kinh), China (Han, Uyghur), Korea
          • B4a – Korea, Han Chinese (Denver), Tujia, Uyghur, Borneo (Bidayuh[19])
            • B4a1 (TMRCA 22,900 [95% CI 18,200 <-> 28,400] ybp[20])
              • B4a1a (TMRCA 9,700 [95% CI 9,000 <-> 10,500] ybp[20])
                • B4a1a* – Philippines (Ivatan, etc.), Malaysia, Papua New Guinea (Trobriand Islands), Ireland
                • B4a1a1 (A14022G, A16247G) – (TMRCA 7,000 [95% CI 6,600 <-> 7,500] ybp[20]) Vanuatu (Port Olry), Papua New Guinea (Siwai of Bougainville)[21] the Polynesian motif,[22][23][24][25] or "PM" (though sometimes referred to as its immediate precursor)
                  • B4a1a1a (16247) – (TMRCA 5,400 [95% CI 4,900 <-> 5,900] ybp[20]) Vanuatu (Banks and Torres), Cook Islands (also sometimes referred to as "the Polynesian motif")
                    • B4a1a1a1 – Solomon Islands (Ranongga, Malaita)
                      • B4a1a1a1a – Solomon Islands (Savo)
                        • B4a1a1a1a1 – Solomon Islands (Gela, Isabel)
                      • B4a1a1a1b – Solomon Islands (Gela, Simbo)
                      • B4a1a1a1c – Papua New Guinea (Nasioi and Nagovisi of Bougainville)
                      • B4a1a1a1d – Tonga
                    • B4a1a1a2 – Solomon Islands (Choiseul), Papua New Guinea (Lihir Island)
                      • B4a1a1a2a – Solomon Islands (Malaita)
                      • B4a1a1a2b – Papua New Guinea (Buin of Bougainville)
                    • B4a1a1a3 – Solomon Islands (Malaita, Makira)
                    • B4a1a1a4 – Papua New Guinea (South Coast), Solomon Islands (Guadalcanal)
                    • B4a1a1a5 – Solomon Islands (Malaita, Ontong Java)
                    • B4a1a1a6 – Solomon Islands (Malaita, Vella Lavella)
                    • B4a1a1a7 – Solomon Islands (Bellona)
                    • B4a1a1a8 – Solomon Islands (Tikopia), Fiji
                    • B4a1a1a9 – Solomon Islands (Tikopia)
                    • B4a1a1a10 – Solomon Islands (Savo, Ranongga)
                    • B4a1a1a11 – Solomon Islands (Simbo)
                      • B4a1a1a11a – Solomon Islands (Choiseul), Vanuatu (Banks and Torres)
                      • B4a1a1a11b – Solomon Islands (Bellona), Cook Islands
                    • B4a1a1a12 – Solomon Islands (Gela, Savo)
                    • B4a1a1a13 – Solomon Islands (Choiseul), Samoa
                    • B4a1a1a14 – Papua New Guinea (Buka)
                    • B4a1a1a15 – Tonga, Wallis and Futuna (Futuna)
                    • B4a1a1a16 – Solomon Islands (Tikopia), Tonga
                    • B4a1a1a17 – Papua New Guinea (Buka, Siwai of Bougainville)
                    • B4a1a1a18 – Cook Islands
                    • B4a1a1a19 – Papua New Guinea (Lihir Island, Anem of New Britain)
                    • B4a1a1a20 – Tuvalu
                    • B4a1a1a21 – Solomon Islands (Malaita), Samoa
                    • B4a1a1a22 – Niue, Samoa
                    • B4a1a1a23 – Papua New Guinea (Torau of Bougainville), Solomon Islands (Isabel, Vella Lavella, Shortlands)
                  • B4a1a1b – Madagascar (Mikea, Merina) (Malagasy motif – a Polynesian motif found only among the Malagasy people)
                  • B4a1a1c – Cook Islands
                  • B4a1a1d – Solomon Islands (Isabel), Papua New Guinea (Kavieng)
                  • B4a1a1e – Solomon Islands (Ranongga, Malaita)
                  • B4a1a1f – Solomon Islands (Guadalcanal)
                  • B4a1a1g – Solomon Islands (Russell, Malaita)
                  • B4a1a1h – Solomon Islands (Bellona, Rennell)
                  • B4a1a1i – Solomon Islands (Ranongga, Savo)
                  • B4a1a1j – Solomon Islands (Russell, Guadalcanal)
                  • B4a1a1k – Tonga, Samoa
                    • B4a1a1k1 – Tonga, Samoa
                  • B4a1a1m – Tonga, Samoa, Wallis and Futuna (Futuna)
                    • B4a1a1m1 – Cook Islands, Tuvalu
                  • B4a1a1n – Solomon Islands (Santa Cruz), Cook Islands
                  • B4a1a1o – Papua New Guinea (Madang), Solomon Islands (Tikopia), Samoa
                  • B4a1a1p – Solomon Islands (Gela)
                  • B4a1a1q – Indonesia (West New Guinea), Solomon Islands (Choiseul)
                  • B4a1a1r – Cook Islands
                  • B4a1a1s – Papua New Guinea (Torau and Nagovisi of Bougainville)
                  • B4a1a1t – Samoa, Cook Islands
                  • B4a1a1u – Fiji, Wallis and Futuna (Futuna)
                  • B4a1a1v – Tonga, Wallis and Futuna (Futuna)
                  • B4a1a1w – Papua New Guinea (Anem of New Britain)
                  • B4a1a1x – Tuvalu, Micronesia (Majuro Atoll)
                  • B4a1a1y – Solomon Islands (Vella Lavella)
                  • B4a1a1z – Papua New Guinea (Nakanai of New Britain)
                  • B4a1a1aa – Bougainville (Torau, etc.)
                  • B4a1a1ab – Solomon Islands (Ontong Java), Samoa
                  • B4a1a1ac – Solomon Islands (Kolombangara), Tuvalu
                  • B4a1a1ad – Wallis and Futuna (Futuna)
                  • B4a1a1ae – Papua New Guinea (Kavieng)
                  • B4a1a1af – Papua New Guinea (Anem of New Britain)
                • B4a1a2 – Taiwan (Amis)
                • B4a1a3 – Taiwan (Ami)
                  • B4a1a3a – Taiwan (Siraya)
                    • B4a1a3a1 – Philippines (Ivatan), Malaysia (Kota Kinabalu), Spain, USA
                      • B4a1a3a1a – Taiwan (Amis)
                • B4a1a4 – Philippines (Ivatan), Orchid Island (Yami)
                • B4a1a5 – Philippines, Malaysia (Kota Kinabalu)
                  • B4a1a5a – Philippines (Kalangoya, Ivatan)
                • B4a1a6 – Philippines (Kalangoya, Ifugao)
                  • B4a1a6a – Philippines (Kalangoya, Ibaloi)
                • B4a1a7 – Taiwan (Amis)
              • B4a1b'e (TMRCA 20,000 [95% CI 15,300 <-> 25,700] ybp[20])
                • B4a1b'e* – China (Naxi, Nyingchi, etc.)
                • B4a1b – Japan
                  • B4a1b1 – Japan
                    • B4a1b1a – Japan, Korea
                • B4a1e – China, Taiwan (Makatao), Vietnam (Thái), Thailand (Khon Mueang in Chiang Mai Province, Lamphun Province, and Lampang Province,[18] Tai Yuan in Northern Thailand[26])
              • B4a1c (TMRCA 20,200 [95% CI 15,600 <-> 25,700] ybp) – India, China (Uyghur), Vietnam (Tay), Japan
                • B4a1c1 (TMRCA 17,400 [95% CI 10,700 <-> 26,600] ybp) – Japan
                  • B4a1c1a (TMRCA 13,800 [95% CI 8,200 <-> 21,800] ybp) – Japan, Korea, China
                    • B4a1c1a1 – Japan, Korea
                • B4a1c2'4'5 (TMRCA 17,100 [95% CI 11,800 <-> 23,900] ybp) – Vietnam (Cờ Lao)
                  • B4a1c2 – Tuvan, Tofalar
                  • B4a1c4 (TMRCA 13,400 [95% CI 11,000 <-> 16,300] ybp) – China (Mongol in Hulun Buir, Dai), Vietnam (Dao, Hà Nhì, Si La, Kinh, Nùng), Thailand (Khon Mueang in Chiang Mai Province, Phutai in Sakon Nakhon Province, Nyaw in Nakhon Phanom Province, Lao Isan in four provinces of Northeast Thailand, Shan in Mae Hong Son Province, Htin in Phayao Province, Phuan in Suphan Buri Province[18])
                  • B4a1c5 – China (Fujian), Taiwan (Hakka)
                • B4a1c3 (TMRCA 16,100 [95% CI 10,100 <-> 24,500] ybp[20])
                  • B4a1c3a (TMRCA 3,600 [95% CI 1,650 <-> 6,800] ybp[20]) – Japan, Korea, Kazakh (Zhan Aul of Altai Republic[27]), Kyrgyz (Kyrgyzstan[28])
                  • B4a1c3b (TMRCA 11,500 [95% CI 5,600 <-> 21,200] ybp[20]) – Japan, Korea, China
              • B4a1d – Vietnam
            • B4a2 – Japan
              • B4a2a – Indonesia (Semende of Sumatra, Banjarmasin), Philippines, Taiwan (Makatao, Hakka)
                • B4a2a1 – Orchid Island (Yami), Philippines (Ivatan)
                • B4a2a2 – Taiwan (Atayal, Saisiat)
                • B4a2a3 – Taiwan (Paiwan, Hakka)
              • B4a2b – China (Han from Beijing)
                • B4a2b1 – China, Jamaica
                  • B4a2b1a – Japan
            • B4a3 – Tibet (Nagqu), Japan
            • B4a4 – Ladakh, Northern Areas of Pakistan (Balti), Singapore, China (Han from Beijing, etc.), Russia, Germany
              • B4a4a - Yakut, Yukaghir
              • B4a4b - China
              • B4a4c - Thailand
                • B4a4c1 - Naxi, Uyghur
              • B4a4d - China
              • B4a4e - China
                • B4a4e1 - China, Taiwan
              • B4a4f - Japan
                • B4a4f1 - China
            • B4a5 – China (Han), Taiwan (Hakka), Vietnam (H'Mông, Dao, Cờ Lao)
          • B4g
            • B4g1 – Thailand
              • B4g1a – Thailand (Khon Mueang in Mae Hong Son, Chiang Rai, and Lampang provinces,[18] Phutai in Sakon Nakhon Province[18]), Vietnam (Thái, Nùng, etc.), China (Han from Zhanjiang, etc.)
              • B4g1b – Han Chinese (Beijing, Denver)
            • B4g2 – Orchid Island (Tao), China (Han from Hunan), Vietnam (Cờ Lao, Dao, Si La), Thailand (Tai Dam in Kanchanaburi Province,[18] Phutai in Sakon Nakhon Province,[18] Lao Isan in Chaiyaphum Province,[18] Htin in Phayao Province[18])
          • B4h – China, Taiwan, Vietnam, Thailand (Phuan in Sukhothai Province[18])
            • B4h1 – China (Fujian, etc.), Taiwan, Thailand (Tai Dam in Kanchanaburi Province[18]), Japan
          • B4i – China
            • B4i1 – China (Han from Beijing, etc.)
          • B4k – China (Han from Beijing, etc.)
          • B4m – Korea, China, Taiwan (Minnan), Vietnam
        • B4b'd'e'j – Vietnam, Laos
          • B4b – Canada
            • B2 – Quechua, Guarani, Coreguaje, Waunana, Katuena, Ache, Gaviao, Xavante, Peru, Ecuador, Colombia, Argentina, USA (Yaqui, Hispanics, etc.), Dominican Republic
              • B2a – Northwestern Canada (Tsimshian), Mexico (Chihuahua)
                • B2a1 – USA (Jemez in New Mexico, Hispanics, etc.), Mexico
                  • B2a1a – USA (Hispanics)
                    • B2a1a1 – Mexico (Chihuahua)
                  • B2a1b – Mexico (Chihuahua), USA (Hispanic)
                • B2a2 – USA (New Mexico, Colorado, Mexican)
                • B2a3 – Mexico (Chihuahua, Durango), USA (Mexican)
                • B2a4
                  • B2a4a – Mexico (Sinaloa)
                    • B2a4a1 – Mexico (Chihuahua, Jalisco, Durango)
                • B2a5 – Pima, USA (Arizona, Utah, California)
              • B2b – Cayapa, Pomo, Xavante, Colombia, Peru, Ecuador, Argentina
                • B2b1 – Venezuela, Ecuador (Shuar of Gualaceo)
                • B2b2 – Bolivia (Beni), Argentina (Criollo of Gran Chaco), USA (Hispanic)
                  • B2b2a – Bolivia (Santa Cruz, Cochabamba)
                • B2b3 – Yanomama
                  • B2b3a – Puerto Rico, Venezuela, Kayapo
                • B2b4 – USA (Mexican)
              • B2c – Ecuador, USA (Hispanic), ancient Canada
                • B2c1 – Mexico (Mixe), USA (Hispanic, Mexican)
                  • B2c1a – USA (Mexican, Hispanic)
                  • B2c1b – USA (Hispanic, Mexican)
                  • B2c1c – USA (Mexican)
                • B2c2 – USA (Mexican)
                  • B2c2a – USA (Mexican, Hispanic)
                  • B2c2b – USA (Mexican, Hispanic)
              • B2d – Nicaragua (Chinandega), Ngöbe/Guaymi, Wayuu, Colombia, USA (Hispanic in New Jersey)
              • B2e – Colombia, Argentina, Waiwai
              • B2f – USA (Mexican)
              • B2g
                • B2g1 – Mexico, USA (Yaqui, Mexican, Hispanic)
                • B2g2
              • B2h – Ache
              • B2i
                • B2i1 – Kayapo
                • B2i2 – Chile
                  • B2i2a – Mapuche
                    • B2i2a1 – Chile
                      • B2i2a1a – Chile, Argentina
                      • B2i2a1b – Chile
                  • B2i2b – Chile
                    • B2i2b1 – Chile
              • B2j
              • B2k – Venezuela, USA (Mexican)
              • B2l – Venezuela, Ecuador
              • B2m
              • B2n
              • B2o – Colombia, Mexico (Maya), USA (Hispanic in Arizona)
                • B2o1 – Ecuador, Bolivia
                  • B2o1a – Colombia, Bolivia, Peru
              • B2p – USA (Mexican)
              • B2q – Ecuador, USA (Mexican)
              • B2r – USA (Hispanic, Mexican)
              • B2s – USA (Mexican)
              • B2t – Guatemala (Maya, la Tinta)
              • B2u
              • B2v
              • B2w
              • B2x
              • B2y – South America (Andes), Peru
                • B2y1 – USA
            • B4b1
              • B4b1* – Thailand (Phuan in Phrae Province),[18] Korea, Japan
              • B4b1a
                • B4b1a* – China, Tubalar,[14] Philippines, Indonesia
                • B4b1a-G207A (TMRCA 14,900 [95% CI 11,200 <-> 19,400] ybp[20]) – Japan
                  • B4b1a1 (TMRCA 3,000 [95% CI 2,100 <-> 4,200] ybp[20]) – Japan
                    • B4b1a1a – Japan, Korea
                    • B4b1a1b – Japan
                    • B4b1a1c – Japan, Korea
                  • B4b1a2 (TMRCA 11,900 [95% CI 10,300 <-> 13,600] ybp[20]) – Japan, Korea, China (Fujian), Taiwan, Philippines (Aeta of Bataan, etc.), Indonesia, Thailand (Khon Mueang in Lampang Province[18]), India
                    • B4b1a2a – Thailand (Khon Mueang in Chiang Mai, Lamphun, and Lampang provinces, Phutai in Sakon Nakhon Province, Tai Dam in Loei Province, Lao Isan in Ubon Ratchathani Province[18]), Vietnam (Gelao), China (Han from Zhanjiang), Japan
                    • B4b1a2b – Taiwan (Ami)
                      • B4b1a2b1 – Philippines (Maranao, Manobo)
                      • B4b1a2b2 – Taiwan (Bunun, Makatao)
                    • B4b1a2c – Philippines (Mamanwa)
                    • B4b1a2d – Philippines (Surigaonon)
                    • B4b1a2e – China (She people, etc.)
                    • B4b1a2f – Taiwan (Bunun, Tsou)
                    • B4b1a2g – Taiwan (Bunun)
                      • B4b1a2g1 – Taiwan (Bunun)
                    • B4b1a2h – Taiwan (Ami)
                    • B4b1a2i
                      • B4b1a2i* – Tuvalu, Banjar (Banjarmasin)[citation needed]
                      • B4b1a2i1
                        • B4b1a2i1*
                        • B4b1a2i1a
                          • B4b1a2i1a* – Nauru, Kiribati
                            • B4b1a2i1a1 – Tuvalu
                      • B4b1a2i2 – Solomon Islands (Guadalcanal)
                  • B4b1a3 (TMRCA 7,300 [95% CI 4,600 <-> 11,000] ybp[20]) – Han Chinese (Denver)
              • B4b1b'c
                • B4b1b – Japan, Korean, China (Lanzhou[31]), Vietnam
                • B4b1c – China, Taiwan, South Korea, Japan (TMRCA 14,900 [95% CI 9,800 <-> 21,700] ybp)
                  • B4b1c1 – Vietnam (Kinh, Tày, Nùng), Thailand, China, Japan (TMRCA 7,200 [95% CI 4,500 <-> 10.800] ybp)
                  • B4b1c2 – Mongol (New Barag Left Banner), China, Taiwan (Hakka), Japan (TMRCA 12,900 [95% CI 7,800 <-> 20,100] ybp)
          • B4d
            • B4d1'2'3
              • B4d1'2'3* – Russia (Buryat[14]), China (Oroqen,[29] Tibetan from Tingri, etc.), Korea
              • B4d1 – China (Miao, Han from Fengcheng, Lanzhou,[31] Jiangsu, etc.), Taiwan, Japan (Chiba), conqueror period Hungary (three specimens from the Karos-III site)[32]
                • B4d1a – Han Chinese (Denver), Barghut (Hulun Buir)[16]
              • B4d2 – China (Han from Qingdao)
              • B4d3 – China (Han from Beijing, etc.)
                • B4d3a – China, Italy (TMRCA 8,300 [95% CI 4,700 <-> 13,500] ybp)
                  • B4d3a1 – Japan (Aichi, Ibaraki, etc.), Korea
            • B4d4 – Japan (Chiba, etc.)
          • B4e – Thailand (Phuan in Lopburi, Sukhothai, and Phrae provinces,[18] Tai Yuan in Uttaradit Province[18]), Laos (Lao in Vientiane[18]), Vietnam (La Hủ), China, Japan (Tokyo)
          • B4j – Buryat, Khamnigan[16]
        • B4c – Thailand, Indonesia
        • B4f – Japan (Japanese, Ryukyuan, Ainu, late 3–4th century AD (early Kofun period) Yokohama[36]), Korea, Vietnam (Lô Lô)
          • B4f* – Vietnam (Lô Lô), Japan (Aichi)
          • B4f1 - Barghut[16]
            • B4f1* – Japan (Tokyo)
            • B4f1a – Japan (Tokyo, etc.)
      • B5
        • B5* – China
        • B5a – Thailand (Tai Dam from Kanchanaburi Province[18]), Vietnam (Kinh), China (Han), Taiwan (Hakka), Philippines (Agta of Iriga)
          • B5a1 – Thailand (Tai Yuan from Ratchaburi Province, Blang from Chiang Rai Province, Lao Isan from Chaiyaphum Province, Nyaw from Nakhon Phanom Province, Tai Dam from Kanchanaburi Province, Phuan from Sukhothai Province, Soa from Sakon Nakhon Province[18]), Indonesia (Besemah of Sumatra), Vietnam, China
            • B5a1a – Cambodia, Vietnam (Kinh, Gelao), Laos,[18] Thailand,[18] Indonesia (Besemah and Kutaradja of Sumatra), China, Uyghur, Taiwan (Minnan), Philippines, India
              • B5a1a1 – Nicobar Islands
            • B5a1b – China (Han from Wuhan), Philippines, Iran[37]
              • B5a1b1 – Cambodia (Jarai), Indonesia, Philippines, Taiwan, Thailand (Bru from Sakon Nakhon Province, Phuan from Sukhothai Province and Lopburi Province, Tai Yuan from Uttaradit Province, Khon Mueang from Mae Hong Son Province and Chiang Mai Province, Tai Dam from Kanchanaburi Province, Soa from Sakon Nakhon Province, Nyaw from Nakhon Phanom Province, Saek from Nakhon Phanom Province[18]), Laos (Lao from Luang Prabang and Vientiane[18]), China, Korea
            • B5a1c – China, Taiwan (Minnan), Thailand (Kaleun from Nakhon Phanom Province[18]), Guyana
              • B5a1c1 – China, Taiwan (Minnan), Laos (Lao from Luang Prabang[18])
                • B5a1c1a – Han Chinese
                  • B5a1c1a1 – China (Han from Hunan, etc.)
              • B5a1c2 – China (Han)
            • B5a1d – China, Thailand, Laos, Cambodia, Indonesia (Semende of Sumatra)
          • B5a2 – China (Han from Hunan)
            • B5a2a
              • B5a2a1 – Han Chinese
                • B5a2a1a – China
                • B5a2a1b – Japan (Tokyo, Chiba), Korea
              • B5a2a2
                • B5a2a2* – China
                • B5a2a2a
                  • B5a2a2a1 – Taiwan (Paiwan, Rukai)
                  • B5a2a2a2 – Taiwan (Saisiyat, Rukai)
                • B5a2a2b
                  • B5a2a2b1 – Philippines (Ivatan)
                    • B5a2a2b1a – Taiwan (Bunun)
                  • B5a2a2b2 – Taiwan (Makatao)
        • B5b – China, Uyghur, Kyrgyz
          • B5b1 – China, Tibet, Buryat (Inner Mongolia), Korea, Japan (Tokyo, etc.), Thailand (Suay from Surin Province[18]), Cambodia (Lao), Vietnam, Singapore
            • B5b1a – China, Thailand (Shan from Mae Hong Son Province[18])
              • B5b1a1 – Japan (Tokyo, Aichi)
              • B5b1a2 – Japan (Tokyo)
                • B5b1a2a – Japan (Chiba, Tokyo)
            • B5b1c – Philippines (Ivatan, etc.), Solomon Islands (Isabel, Santa Cruz), Malaysia (Jawa, Batek), Singapore, Yemen
              • B5b1c1 – Philippines (Kalangoya, Ifugao, Ibaloi, Kankanaey)
                • B5b1c1a – Philippines (Kankanaey, Ifugao, Kalangoya, Ibaloi, Abaknon)
          • B5b2 – Russia (Russian old settler in Pokhosk Village of Sakha Republic, Ulchi,[38] Altaian Kazakh[16]), China (Han, Uyghur, Barghut[16]), Japan, Philippines
            • B5b2a – Negidal, Khamnigan
              • B5b2a1 – Japan (Aichi, Tokyo, etc.), China (Han from Wuhan)
              • B5b2a2 – Japan (Tokyo, etc.), Buryat, Hezhen
                • B5b2a2a
                  • B5b2a2a1 – Japan (Tokyo, Chiba, Aichi)
                  • B5b2a2a2 – Malaysia (Bidayuh of Sarawak),[19] Philippines, Solomon Islands (Ranongga)
            • B5b2b – Yakut
            • B5b2-C204T! - China (Han), Korea, Vietnam (Kinh)
              • B5b2c – Taiwan (Minnan, Makatao)
                • B5b2c1 – China (Han from Hunan), Japan (Chiba, Aichi)
          • B5b3
          • B5b4 – China, Altai Kizhi
          • B5b5 – Taiwan (Hakka), Han Chinese (Denver)
    • R11'B6
      • R11 – China (Han from Beijing)
      • B6
        • B6a – China (Han from Tai'an), Thailand (Htin in Phayao Province, Palaung and Khon Mueang in Chiang Mai Province, Phuan in Phrae Province and Sukhothai Province, Mon in Ratchaburi Province and Lopburi Province, Tai Dam in Kanchanaburi Province[18])
          • B6a1 – China, Thailand (Khon Mueang in Lampang Province, Htin in Phayao Province, Blang in Chiang Rai Province[18]), Philippines
            • B6a1a – Myanmar, Thailand (Lawa and Shan in Mae Hong Son Province, Khon Mueang in Chiang Mai Province[18]), Malaysia (Temuan)
    • R24 – Philippines (Mamanwa)
      • R24a – Philippines

Popular culture[edit]

In his popular book The Seven Daughters of Eve, Bryan Sykes named the originator of this mtDNA haplogroup Ina.

See also[edit]

Phylogenetic tree of human mitochondrial DNA (mtDNA) haplogroups

  Mitochondrial Eve (L)    
L0 L1–6  
L1 L2   L3     L4 L5 L6
M N  
CZ D E G Q   O A S R   I W X Y
C Z B F R0   pre-JT   P   U
HV JT K
H V J T

References[edit]

  1. ^ a b van Oven, Mannis; Manfred Kayser (13 Oct 2008). "Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation". Human Mutation. 30 (2): E386–E394. doi:10.1002/humu.20921. PMID 18853457. S2CID 27566749.
  2. ^ Yong-Gang Yao et al. 2001, Phylogeographic Differentiation of Mitochondrial DNA in Han Chinese Am J Hum Genet. 2002 March; 70(3): 635–651
  3. ^ "Downloadable genotypes of present-day and ancient DNA data (compiled from published papers) | David Reich Lab". reich.hms.harvard.edu. Archived from the original on 2019-11-02. Retrieved 2019-09-11.
  4. ^ Haplogroup B.
  5. ^ Fagundes, Nelson J.R.; Ricardo Kanitz; Roberta Eckert; Ana C.S. Valls; Mauricio R. Bogo; Francisco M. Salzano; David Glenn Smith; Wilson A. Silva; Marco A. Zago; Andrea K. Ribeiro-dos-Santos; Sidney E.B. Santos; Maria Luiza Petzl-Erler; Sandro L.Bonatto (2008). "Mitochondrial Population Genomics Supports a Single Pre-Clovis Origin with a Coastal Route for the Peopling of the Americas" (PDF). American Journal of Human Genetics. 82 (3): 583–592. doi:10.1016/j.ajhg.2007.11.013. PMC 2427228. PMID 18313026. Archived from the original (PDF) on 2009-03-25. Retrieved 2009-11-19.
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  11. ^ a b c d e Kristina A. Tabbada, Jean Trejaut, Jun-Hun Loo et al., "Philippine Mitochondrial DNA Diversity: A Populated Viaduct between Taiwan and Indonesia?" Mol. Biol. Evol. 27(1):21–31. (2010) doi:10.1093/molbev/msp215
  12. ^ Min-Sheng Peng, Jun-Dong He, Hai-Xin Liu, and Ya-Ping Zhang, "Tracing the legacy of the early Hainan Islanders – a perspective from mitochondrial DNA," BMC Evol Biol. 2011; 11: 46. Published online 2011 February 15. doi: 10.1186/1471-2148-11-46
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