Haplogroup B (mtDNA)

Haplogroup B
Possible time of origin 50,000 YBP
Possible place of origin Southeastern Asia
Ancestor R11'B
Descendants B4, B5, B7
Defining mutations 8281-8289d[1]

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

Origin

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

Its greater variety is in China. It is conspicuous that haplogroup B may have its earliest diversification in southern China and/or Southeast Asia.[2]

Distribution

Haplogroup B is found frequently in southeastern Asia.[3] A subclade of B4b that is labelled irregularly as B2 is one of five haplogroups found in 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.[4] However, Haplogroup B has been found among Turkic, Mongolic, and Tungusic populations of Siberia, such as Tuvans, Altays, Shors, Khakassians, Yakuts, Buryats, Khamnigans, Negidals, and Evenks.[5] This haplogroup is also found among populations in China, Indonesia, Iran,[5] Iraq,[6] Japan, Korea, Laos,[7] Madagascar, Malaysia, Melanesia, Micronesia, Mongolia, the Philippines, Polynesia, Taiwan, Thailand, Tibet, and Vietnam.[8][9]

Although haplogroup B in general has been found in many samples of Altaic-speaking populations of Siberia, 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%).[10][11][12] 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),[13] Miyazaki and Tokyo, Japan (approx. 3%),[14] South Korea (4/185 = 2.2%),[9] Tuva (1/95 = 1.1% Tuvan),[13] and Hulunbuir (1/149 = 0.7% Barghut).[15]

Table of Frequencies of MtDNA Haplogroup B

Population Frequency Count Source Subtypes
Carolinian (Saipan)1.00017Vilar 2013B4a1a1a=15, B4a1a(xB4a1a1a)=2
Dingban Yao (Mengla, Yunnan)0.60010Wen 2005B4(xB4a, B4b1, B4e, B4f)=3, B4a=1, B4e=1, B4f=1
Iu Mien (Mengla, Yunnan)0.48127Wen 2005B4a=7, B4b1=5, B5a=1
Bapai Yao (Liannan, Guangdong)0.42935Wen 2005B5a=7, B4e=3, B4b1=2, B4a=1, B4(xB4a, B4b1, B4e, B4f)=1, B(xB4, B5)=1
Filipino0.42264Tabbada 2010B4a1a(xB4a1a1)=8, B5b=7, B4b1=6, B4c1b=4, B4a1a1=1, B5a=1
Tu Yao (Hezhou, Guangxi)0.39041Wen 2005B5a=7, B4(xB4a, B4b1, B4e, B4f)=6, B4a=3
Guoshan Yao (Jianghua, Hunan)0.37524Wen 2005B5a=7, B4(xB4a, B4b1, B4e, B4f)=2
Cham (Bình Thuận, Vietnam)0.369168Peng 2010B5a=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)0.36819Wen 2005B5a=3, B4a=2, B4b1=1, B4e=1
Filipino (Luzon)0.367177Tabbada 2010B4a1a(xB4a1a1)=20, B5b=18, B4b1=13, B4c1b=10, B5a=2, B4a(xB4a1a)=1, B7=1
Hmong (Wenshan, Yunnan)0.33339Wen 2005B5a=8, B4a=5
Dong (Tianzhu County, Guizhou)0.32128Li 2007B4a=6, B5a=2, B4c=1
Kinh (Vietnamese)0.317139He 2012B4=24, B5=19, B6=1
Mien (Shangsi, Guangxi)0.31332Wen 2005B4a=5, B5a=3, B4e=2
Kim Mun (Malipo, Yunnan)0.30040Wen 2005B5a=5, B4a=3, B4e=2, B4b1=1, B5(xB5a, B5b)=1
Dai (Xishuangbanna, Yunnan)0.29341Yao 2002B5a=4, B(xB4, B5a, B5b)=4, B4a=3, B4(xB4a)=1
Bunu (Dahua & Tianlin, Guangxi)0.28025Wen 2005B4a=5, B5a=2
Filipino (Visayas)0.277112Tabbada 2010B4a1a(xB4a1a1)=11, B4b1=8, B4c1b=7, B5b=5
Lanten Yao (Tianlin, Guangxi)0.26926Wen 2005B4a=3, B5a=2, B4b1=1, B4f=1
Laos0.262214Bodner 2011B5a=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)0.25831Wen 2005B4a=4, B5a=3, B4b1=1
Lahu (Simao, Yunnan)0.25032Wen 2004B4a=5, B4(xB4a, B4b1)=3
Pan Yao (Tianlin, Guangxi)0.25032Wen 2005B5a=4, B4a=2, B4f=1, B5b=1
Yi (Xishuangbanna, Yunnan)0.25016Wen 2004B4(xB4a, B4b1)=1, B4a=1, B5(xB5a, B5b)=1, B5a=1
Naxi (Lijiang, Yunnan)0.24445Wen 2004B4a=5, B4(xB4a, B4b1)=3, B5a=3
Filipino (Mindanao)0.24370Tabbada 2010B4a1a(xB4a1a1)=7, B4b1=5, B5b=3, B4a(xB4a1a)=1, B4a1a1=1
Lowland Yao (Fuchuan, Guangxi)0.23842Wen 2005B4a=3, B4b1=3, B4(xB4a, B4b1, B4e, B4f)=2, B5a=1, B5b=1
Han (Taiwanese)0.234111Chen 2013B4+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)0.22631Li 2007B4a=3, B4(xB4a, B4b, B4c)=2, B4b=1, B5a=1
Bai (Xishuangbanna, Yunnan)0.21119Wen 2004B4(xB4a, B4b1)=2, B4a=1, B5a=1
Taiwanese (Taipei, Taiwan)0.20991Umetsu 2005B4(xB4f)=13, B5=6
Gelao (Daozhen County, Guizhou)0.206102Liu 2011B4a=5, B4b1=5, B5a=5, B4(xB4a, B4b, B4c, B4e, B4f, B4g)=3, B5b=2, B4c=1
Korean (South Korea)0.204103Derenko 2007B4=13, B5=8
CHS (Han from Hunan & Fujian)0.20055Zheng 2011B5=6, B4=5
Lahu (Xishuangbanna, Yunnan)0.20015Wen 2004B4(xB4a, B4b1)=1, B4a=1, B4b1=1
Han (Southern California)0.192390Ji 2012B=75
Dai (Xishuangbanna, Yunnan)0.19021Qian 2001B4a=2, B4(xB4a)=1, B5b=1
Han (Southwest China; pool of 44 Sichuan, 34 Chongqing, 33 Yunnan, and 26 Guizhou)0.190137Ji 2012B4b'd=9, B4a=6, B5a=4, B4c=3, B(xB4a, B4b'd, B4c, B5a, B5b, B6)=2, B5b=1, B6=1
Tujia (western Hunan)0.18864Wen 2004B4a=5, B5b=3, B4(xB4a, B4b1)=2, B5a=2
Hmong (Jishou, Hunan)0.184103Wen 2005B5a=6, B4a=5, B4b1=3, B4(xB4a, B4b1, B4e, B4f)=2, B4e=1, B4f=1, B5b=1
Hani (Xishuangbanna, Yunnan)0.18233Wen 2004B5a=2, B4(xB4a, B4b1)=1, B4a=1, B4b1=1, B5b=1
Xiban Yao (Fangcheng, Guangxi)0.18211Wen 2005B4a=1, B4b1=1
Korean (Seoul & Daejeon, South Korea)0.180261Kim 2008B(xB5)=32, B5=15
Han (Taiwan)0.1791117Ji 2012B=200
CHD (Han from Denver, Colorado)0.17873Zheng 2011B4=12, B5=1
Hui (Xinjiang)0.17845Yao 2004B4a=4, B5a=2, B4b1=1, B6=1
Yi (Shuangbai, Yunnan)0.17540Wen 2004B4(xB4a, B4b1)=5, B4a=1, B4b1=1
Tujia (Yanhe County, Guizhou)0.17229Li 2007B4b=3, B5a=2
Thailand0.171105Kaewsutthi 2011B(xB5a1)=13, B5a1=5
Tibetan (Zhongdian, Yunnan)0.17135Wen 2004B5a=4, B4(xB4a, B4b1)=2
Aini (Xishuangbanna, Yunnan)0.16050Wen 2004B4a=4, B(xB4, B5)=2, B4(xB4a, B4b1)=1, B5a=1
Chinese (Shenyang, Liaoning)0.156160Umetsu 2005B4(xB4f)=15, B5=10
Daur (Evenk Autonomous Banner)0.15645Kong 2003B4(xB4a, B4b)=4, B4a=3
Korean (Seoul National University Hospital)0.155633Fuku 2007B=98
Yi (Hezhang County, Guizhou)0.15020Li 2007B4b=1, B4(xB4a, B4b, B4c)=1, B6=1
Mongolian (Ulan Bator)0.14947Derenko 2007B4=5, B5=2
Korean (South Korea)0.148203Umetsu 2005B4(xB4f)=24, B5=6
Korean (South Korea)0.146185Jin 2009B4a=11, B4(xB4a, B4b, B4c)=7, B4b1=4, B5a=2, B5b=2, B4c=1
Vietnamese0.14342Jin 2009B4(xB4a, B4b, B4c)=3, B4a=1, B4b(xB4b1)=1, B5a=1
Okinawa0.141326Umetsu 2005B4(xB4f)=28, B4f=10, B5=8
CHB (Han from Beijing Normal University)0.140121Zheng 2011B4=14, B5=3
JPT (Japanese from Tokyo)0.136118Zheng 2011B4=11, B5=5
Tujia (Yongshun, Hunan)0.13330Wen 2004B4(xB4a, B4b1)=2, B5a=2
Japanese (Hokkaidō)0.129217Asari 2007B4(xB4f)=18, B4f=5, B5=5
Japanese (Tōkai)0.124282Umetsu 2005B4(xB4f)=26, B5=9
Japanese (Gifu)0.1211617Fuku 2007B=196
Negidal0.12133Starikovskaya 2005B5b2=4
Japanese (Tōhoku)0.119336Umetsu 2005B4(xB4f)=24, B5=15, B4f=1
Korean (northern China)0.11851Jin 2009B4a=2, B4b(xB4b1)=2, B5a=1, B5b=1
Telengit (Altai Republic)0.11371Derenko 2007B4=8
Jino (Xishuangbanna, Yunnan)0.11118Wen 2004B4a=1, B5a=1
Tibetan (Nyingchi, Tibet)0.11154Ji 2012B4a=5, B5b=1
Japanese (northern Kyūshū)0.109256Umetsu 2005B4(xB4f)=19, B5=8, B4f=1
Japanese0.109211Maruyama 2003B4b1=14, B5b=4, B5a=2, B4a=1, B4(xB4a, B4b1)=1, B5(xB5a, B5b)=1
Evenk (New Barag Left Banner)0.10647Kong 2003B4b=4, B5b=1
Korean (Arun Banner)0.10448Kong 2003B4b=2, B5b=2, B4(xB4a, B4b)=1
Mongol (New Barag Left Banner)0.10448Kong 2003B4b=3, B4a=1, B5a=1
Han (Beijing)0.10040Jin 2009B(xB4, B5a, B5b)=2, B4b(xB4b1)=2
Nu (Gongshan, Yunnan)0.10030Wen 2004B4(xB4a, B4b1)=3
Thai0.10040Jin 2009B5a=3, B(xB4, B5a, B5b)=1
Yi (Luxi, Yunnan)0.09731Wen 2004B4(xB4a, B4b1)=2, B(xB4, B5)=1
Japanese (Miyazaki)0.090100Uchiyama 2007B4b1a1=3, B4(xB4a, B4b1)=2, B4a=2, B5b=2
Khakassian (Khakassia)0.08857Derenko 2007B4=5
Han (Xinjiang)0.08547Yao 2004B5a=2, B4b1=1, B5b=1
Kyrgyz (Sary-Tash, Kyrgyzstan)0.08547Yao 2004B5a=2, B4(xB4a, B4b1, B4c)=1, B4a=1
Mongolian (Ulan Bator)0.08547Jin 2009B4(xB4a, B4b, B4c)=2, B4a=1, B5b=1
Tuvan0.08495Starikovskaya 2005B4a1c2=4, B5=2, B4(xB4a1c2, B4b1a, B4d1a)=1, B4b1a=1
Tibetan (Nyingchi, Tibet)0.08324Ji 2012B=2
Barghut (Hulunbuir)0.081149Derenko 2012B4c1a2(xB4c1a2a)=4, B4f1=2, B5b(xB5b2)=2, B4c1a2a=1, B4b1a3a1a=1, B4d1=1, B5b2=1
Chamorro (85 Guam, 14 Saipan, & 6 Rota)0.076105Vilar 2013B4a1a1a=7, B4b1=1
Manchurian0.07540Jin 2009B4(xB4a, B4b, B4c)=2, B4a=1
Uzbek (Uzbekistan/Kyrgyzstan)0.07540Comas 2004B=3
Uyghur (Penjim, Panfilov District, Almaty Province, Kazakhstan)0.07355Yao 2004B5a=3, B4(xB4a, B4b1, B4c)=1
Buryat0.071126Kong 2003B4(xB4a, B4b)=5, B5b=2, B(xB4, B5a, B5b)=1, B4b=1
Khamnigan (Buryatia)0.07199Derenko 2007B4=5, B5=2
Dungan (Uzbekistan/Kyrgyzstan)0.06316Comas 2004B=1
Persian (eastern Iran)0.06182Derenko 2007B4=4, B5=1
Nogai (Dagestan)0.06133Marchani 2008B=2
Tibetan (Nagchu, Tibet)0.05735Ji 2012B=2
Kazakh (Kegen Valley, Almaty Province, Kazakhstan)0.05555Yao 2004B4(xB4a, B4b1, B4c)=1, B4a=1, B5b=1
Lisu (Gongshan, Yunnan)0.05437Wen 2004B4(xB4a, B4b1)=1, B5(xB5a, B5b)=1
Tharu (Chitwan, Nepal)0.053133Fornarino 2009B5a=7
Uzbek (Xinjiang)0.05258Yao 2004B4b1=2, B4c=1
Bukharan Arab (Uzbekistan/Kyrgyzstan)0.05020Comas 2004B=1
Filipino (Palawan)0.05020Scholes 2011B4a1a=1
Tajik (Uzbekistan/Kyrgyzstan)0.05020Comas 2004B=1
Turkmen (Uzbekistan/Kyrgyzstan)0.05020Comas 2004B=1
Shor (Kemerovo Oblast)0.04982Derenko 2007B4=4
Va (Simao, Yunnan)0.04522Qian 2001B4(xB4a)=1
Altai Kizhi0.04490Derenko 2007B4=3, B5=1
Bai (Dali, Yunnan)0.04468Wen 2004B5a=2, B4b1=1
Tofalar0.04346Starikovskaya 2005B4a1c2=2
Kyrgyz (Bakay-Ata, Kyrgyzstan)0.04248Yao 2004B4(xB4a, B4b1, B4c)=1, B5b=1
Tubalar0.04272Starikovskaya 2005B4b1a=3
Evenk (Krasnoyarsk Krai)0.04173Derenko 2007B4=3
Kazakh (Kosh-Agach, Altai Republic)0.04198Derenko 2012B5b2=3, B4c1b=1
Buryat (Kushun, Nizhneudinsky, Irkutsk Oblast)0.04025Starikovskaya 2005B4d1a=1
Kazakh (Xinjiang)0.03853Yao 2004B4a=2
Teleut (Kemerovo Oblast)0.03853Derenko 2007B4=2
Kalmyk (Kalmykia)0.036110Derenko 2007B4=3, B5=1
Tibetan (Chamdo, Tibet)0.03429Ji 2012B4a=1
Buryat (Buryatia)0.034295Derenko 2007B4=9, B5=1
Lahu (Lancang, Yunnan)0.02935Wen 2004B4(xB4a, B4b1)=1
Pumi (Ninglang, Yunnan)0.02836Wen 2004B4(xB4a, B4b1)=1
Va (Ximeng & Gengma, Yunnan)0.02836Yao 2002B5a=1
Yakut (Yakutia)0.02836Derenko 2007B4=1
Tharu (Morang, Nepal)0.02540Fornarino 2009B5a=1
Tibetan (Deqin, Yunnan)0.02540Wen 2004B4(xB4a, B4b1)=1
Oroqen (Oroqen Autonomous Banner)0.02344Kong 2003B4(xB4a, B4b)=1
Uyghur (Xinjiang)0.02147Yao 2004B(xB4, B5, B6)=1
Oirat Mongol (Xinjiang)0.02049Yao 2004B4b1=1
Ainu0.02051Sato 2009
Tajima 2004
B4f1=1
Tuvinian (Tuva)0.019105Derenko 2007B4=2
Tibetan (Qinghai)0.01856Wen 2004B4a=1
Nogai (Nogaysky, Dagestan & Adyge-Khabalsky, Karachay-Cherkessia)0.015206Bermisheva 2004B=3
Tibetan (Shannan, Tibet)0.01474Ji 2012B4a=1
Bashkir0.009221Bermisheva 2002B=2
Chukchi (Anadyr)0.00015Derenko 2007 -
Uyghur (Uzbekistan/Kyrgyzstan)0.00016Comas 2004 -
Crimean Tatar (Uzbekistan/Kyrgyzstan)0.00020Comas 2004 -
Iranian (Uzbekistan/Kyrgyzstan)0.00020Comas 2004 -
Karakalpak (Uzbekistan/Kyrgyzstan)0.00020Comas 2004 -
Kazakh (Uzbekistan/Kyrgyzstan)0.00020Comas 2004 -
Kyrgyz (Uzbekistan/Kyrgyzstan)0.00020Comas 2004 -
Hindu (Chitwan, Nepal)0.00024Fornarino 2009 -
Nganasan0.00024Starikovskaya 2005 -
Tibetan (Diqing, Yunnan)0.00024Wen 2004 -
Kurd (northwestern Iran)0.00025Derenko 2007 -
Andhra Pradesh (tribal)0.00029Fornarino 2009 -
Tibetan (Shigatse, Tibet)0.00029Ji 2012 -
Batak (Palawan)0.00031Scholes 2011 -
Ket0.00038Starikovskaya 2005 -
Tajik (Tajikistan)0.00044Derenko 2007 -
Tibetan (Lhasa, Tibet)0.00044Ji 2012 -
Evenk (Buryatia)0.00045Derenko 2007 -
Udege (Gvasiugi, Imeni Lazo, Khabarovsk Krai)0.00046Starikovskaya 2005 -
Itelmen0.00047Starikovskaya 2005 -
Chuvash0.00055Bermisheva 2002 -
Nivkh (northern Sakhalin)0.00056Starikovskaya 2005 -
Komi-Zyryans0.00062Bermisheva 2002 -
Mansi0.00063Pimenoff 2008 -
Chukchi0.00066Starikovskaya 2005 -
Evenk (53 Stony Tunguska basin & 18 Tuguro-Chumikan)0.00071Starikovskaya 2005 -
Tatar (Aznakayevo)0.00071Malyarchuk 2010 -
Komi-Permyaks0.00074Bermisheva 2002 -
Siberian Eskimo0.00079Starikovskaya 2005 -
Ulchi0.00087Starikovskaya 2005 -
Mansi0.00098Starikovskaya 2005 -
Udmurt0.000101Bermisheva 2002 -
Mordvinian0.000102Bermisheva 2002 -
Khanty0.000106Pimenoff 2008 -
Yakut0.000117Kong 2003 -
Tatar (Buinsk)0.000126Malyarchuk 2010 -
Mari0.000136Bermisheva 2002 -
Koryak0.000155Starikovskaya 2005 -
Tatar0.000228Bermisheva 2002 -

Subclades

Tree

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.

Popular culture

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

See also

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

  1. 1 2 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.
  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. Haplogroup B.
  4. 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 2427228Freely accessible. PMID 18313026. Retrieved 2009-11-19.
  5. 1 2 Miroslava Derenko, Boris Malyarchuk, Tomasz Grzybowski et al., "Phylogeographic Analysis of Mitochondrial DNA in Northern Asian Populations", Am. J. Hum. Genet. 2007;81:1025–1041. DOI: 10.1086/522933
  6. Nadia Al-Zahery, Maria Pala, Vincenza Battaglia et al., BMC Evolutionary Biology 2011, 11:288. http://www.biomedcentral.com/1471-2148/11/288
  7. Martin Bodner, Bettina Zimmermann, Alexander Röck, Anita Kloss-Brandstätter, David Horst, Basil Horst, Sourideth Sengchanh, Torpong Sanguansermsri, Jürgen Horst, Tanja Krämer, Peter M Schneider, and Walther Parson, "Southeast Asian diversity: first insights into the complex mtDNA structure of Laos." BMC Evolutionary Biology (2011), 11:49. http://www.biomedcentral.com/1471-2148/11/49
  8. Theodore G Schurr and Douglas C Wallace, "Mitochondrial DNA diversity in Southeast Asian populations", Human Biology, June 2002.
  9. 1 2 Jin H-J, Tyler-Smith C, Kim W (2009), "The Peopling of Korea Revealed by Analyses of Mitochondrial DNA and Y-Chromosomal Markers." PLoS ONE 4(1): e4210. doi:10.1371/journal.pone.0004210
  10. 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
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