Haplogroup B (mtDNA)

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Haplogroup B
Peopling of eurasia.jpg
Possible time of origin50,000 YBP
Possible place of origin Southeast Asia
Ancestor R11'B B4'5'R11B6'R24
DescendantsB2,B4, B5, B6
Defining mutations8281-8289d [1]

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

Contents

Origin

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

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

PopulationFrequencyCountSourceSubtypes
Carolinian (Saipan)1.00017[ citation needed ]B4a1a1a=15, B4a1a(xB4a1a1a)=2
Hawaiians 0.994160 [17] B4a1a1, B4a1a1a1, B4a1a1a3
Dingban Yao (Mengla, Yunnan)0.60010[ citation needed ]B4(xB4a, B4b1, B4e, B4f)=3, B4a=1, B4e=1, B4f=1
Iu Mien (Mengla, Yunnan)0.48127[ citation needed ]B4a=7, B4b1=5, B5a=1
Bapai Yao (Liannan, Guangdong)0.42935[ citation needed ]B5a=7, B4e=3, B4b1=2, B4a=1, B4(xB4a, B4b1, B4e, B4f)=1, B(xB4, B5)=1
Filipino 0.42264 [11] B4a1a(xB4a1a1)=8, B5b=7, B4b1=6, B4c1b=4, B4a1a1=1, B5a=1
Tu Yao (Hezhou, Guangxi)0.39041[ citation needed ]B5a=7, B4(xB4a, B4b1, B4e, B4f)=6, B4a=3
Guoshan Yao (Jianghua, Hunan)0.37524[ citation needed ]B5a=7, B4(xB4a, B4b1, B4e, B4f)=2
Cham (Bình Thuận, Vietnam)0.369168 [18] 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)0.36819[ citation needed ]B5a=3, B4a=2, B4b1=1, B4e=1
Filipino (Luzon)0.367177 [11] B4a1a(xB4a1a1)=20, B5b=18, B4b1=13, B4c1b=10, B5a=2, B4a(xB4a1a)=1, B7=1
Hmong (Wenshan, Yunnan)0.33339[ citation needed ]B5a=8, B4a=5
Dong (Tianzhu County, Guizhou)0.32128[ citation needed ]B4a=6, B5a=2, B4c=1
Kinh (Vietnamese)0.317139[ citation needed ]B4=24, B5=19, B6=1
Mien (Shangsi, Guangxi)0.31332[ citation needed ]B4a=5, B5a=3, B4e=2
Kim Mun (Malipo, Yunnan)0.30040[ citation needed ]B5a=5, B4a=3, B4e=2, B4b1=1, B5(xB5a, B5b)=1
Dai (Xishuangbanna, Yunnan)0.29341[ citation needed ]B5a=4, B(xB4, B5a, B5b)=4, B4a=3, B4(xB4a)=1
Bunu (Dahua & Tianlin, Guangxi)0.28025[ citation needed ]B4a=5, B5a=2
Filipino (Visayas)0.277112 [11] B4a1a(xB4a1a1)=11, B4b1=8, B4c1b=7, B5b=5
Lanten Yao (Tianlin, Guangxi)0.26926[ citation needed ]B4a=3, B5a=2, B4b1=1, B4f=1
Laos 0.262214 [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)0.25831[ citation needed ]B4a=4, B5a=3, B4b1=1
Lahu (Simao, Yunnan)0.25032[ citation needed ]B4a=5, B4(xB4a, B4b1)=3
Pan Yao (Tianlin, Guangxi)0.25032[ citation needed ]B5a=4, B4a=2, B4f=1, B5b=1
Yi (Xishuangbanna, Yunnan)0.25016[ citation needed ]B4(xB4a, B4b1)=1, B4a=1, B5(xB5a, B5b)=1, B5a=1
Naxi (Lijiang, Yunnan)0.24445[ citation needed ]B4a=5, B4(xB4a, B4b1)=3, B5a=3
Filipino (Mindanao)0.24370 [11] B4a1a(xB4a1a1)=7, B4b1=5, B5b=3, B4a(xB4a1a)=1, B4a1a1=1
Lowland Yao (Fuchuan, Guangxi)0.23842[ citation needed ]B4a=3, B4b1=3, B4(xB4a, B4b1, B4e, B4f)=2, B5a=1, B5b=1
Han (Taiwan)0.234111[ 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)0.22631[ citation needed ]B4a=3, B4(xB4a, B4b, B4c)=2, B4b=1, B5a=1
Bai (Xishuangbanna, Yunnan)0.21119[ citation needed ]B4(xB4a, B4b1)=2, B4a=1, B5a=1
Han (Taipei, Taiwan)0.20991[ citation needed ]B4(xB4f)=13, B5=6
Gelao (Daozhen County, Guizhou)0.206102[ citation needed ]B4a=5, B4b1=5, B5a=5, B4(xB4a, B4b, B4c, B4e, B4f, B4g)=3, B5b=2, B4c=1
Korean (South Korea)0.204103 [6] B4=13, B5=8
CHS (Han from Hunan & Fujian)0.20055[ citation needed ]B5=6, B4=5
Lahu (Xishuangbanna, Yunnan)0.20015[ citation needed ]B4(xB4a, B4b1)=1, B4a=1, B4b1=1
Han (Southern California)0.192390[ citation needed ]B=75
Dai (Xishuangbanna, Yunnan)0.19021[ citation needed ]B4a=2, B4(xB4a)=1, B5b=1
Han (Southwest China; pool of 44 Sichuan, 34 Chongqing, 33 Yunnan, and 26 Guizhou)0.190137[ 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)0.18864[ citation needed ]B4a=5, B5b=3, B4(xB4a, B4b1)=2, B5a=2
Hmong (Jishou, Hunan)0.184103[ citation needed ]B5a=6, B4a=5, B4b1=3, B4(xB4a, B4b1, B4e, B4f)=2, B4e=1, B4f=1, B5b=1
Hani (Xishuangbanna, Yunnan)0.18233[ citation needed ]B5a=2, B4(xB4a, B4b1)=1, B4a=1, B4b1=1, B5b=1
Xiban Yao (Fangcheng, Guangxi)0.18211[ citation needed ]B4a=1, B4b1=1
Korean (Seoul & Daejeon, South Korea)0.180261[ citation needed ]B(xB5)=32, B5=15
Han (Taiwan)0.1791117[ citation needed ]B=200
CHD (Han from Denver, Colorado)0.17873[ citation needed ]B4=12, B5=1
Hui (Xinjiang)0.17845[ citation needed ]B4a=4, B5a=2, B4b1=1, B6=1
Yi (Shuangbai, Yunnan)0.17540[ citation needed ]B4(xB4a, B4b1)=5, B4a=1, B4b1=1
Tujia (Yanhe County, Guizhou)0.17229[ citation needed ]B4b=3, B5a=2
Thailand 0.171105[ citation needed ]B(xB5a1)=13, B5a1=5
Tibetan (Zhongdian, Yunnan)0.17135[ citation needed ]B5a=4, B4(xB4a, B4b1)=2
Aini (Xishuangbanna, Yunnan)0.16050[ citation needed ]B4a=4, B(xB4, B5)=2, B4(xB4a, B4b1)=1, B5a=1
Chinese (Shenyang, Liaoning)0.156160[ citation needed ]B4(xB4f)=15, B5=10
Daur (Evenk Autonomous Banner)0.15645[ citation needed ]B4(xB4a, B4b)=4, B4a=3
Korean (Seoul National University Hospital)0.155633[ citation needed ]B=98
Yi (Hezhang County, Guizhou)0.15020[ citation needed ]B4b=1, B4(xB4a, B4b, B4c)=1, B6=1
Mongolian (Ulan Bator)0.14947 [6] B4=5, B5=2
Korean (South Korea)0.148203[ citation needed ]B4(xB4f)=24, B5=6
Korean (South Korea)0.146185 [10] B4a=11, B4(xB4a, B4b, B4c)=7, B4b1=4, B5a=2, B5b=2, B4c=1
Vietnamese 0.14342 [10] B4(xB4a, B4b, B4c)=3, B4a=1, B4b(xB4b1)=1, B5a=1
Okinawa 0.141326[ citation needed ]B4(xB4f)=28, B4f=10, B5=8
CHB (Han from Beijing Normal University)0.140121[ citation needed ]B4=14, B5=3
JPT (Japanese from Tokyo)0.136118[ citation needed ]B4=11, B5=5
Tujia (Yongshun, Hunan)0.13330[ citation needed ]B4(xB4a, B4b1)=2, B5a=2
Japanese (Hokkaidō)0.129217[ citation needed ]B4(xB4f)=18, B4f=5, B5=5
Japanese (Tōkai)0.124282[ citation needed ]B4(xB4f)=26, B5=9
Japanese (Gifu)0.1211617[ citation needed ]B=196
Negidal 0.12133 [14] B5b2=4
Japanese (Tōhoku)0.119336[ citation needed ]B4(xB4f)=24, B5=15, B4f=1
Korean (northern China)0.11851 [10] B4a=2, B4b(xB4b1)=2, B5a=1, B5b=1
Telengit (Altai Republic)0.11371 [6] B4=8
Jino (Xishuangbanna, Yunnan)0.11118[ citation needed ]B4a=1, B5a=1
Tibetan (Nyingchi, Tibet)0.11154[ citation needed ]B4a=5, B5b=1
Japanese (northern Kyūshū)0.109256[ citation needed ]B4(xB4f)=19, B5=8, B4f=1
Japanese 0.109211[ citation needed ]B4b1=14, B5b=4, B5a=2, B4a=1, B4(xB4a, B4b1)=1, B5(xB5a, B5b)=1
Evenk (New Barag Left Banner)0.10647[ citation needed ]B4b=4, B5b=1
Korean (Arun Banner)0.10448[ citation needed ]B4b=2, B5b=2, B4(xB4a, B4b)=1
Mongol (New Barag Left Banner)0.10448[ citation needed ]B4b=3, B4a=1, B5a=1
Han (Beijing)0.10040 [10] B(xB4, B5a, B5b)=2, B4b(xB4b1)=2
Nu (Gongshan, Yunnan)0.10030[ citation needed ]B4(xB4a, B4b1)=3
Thai 0.10040 [10] B5a=3, B(xB4, B5a, B5b)=1
Yi (Luxi, Yunnan)0.09731[ citation needed ]B4(xB4a, B4b1)=2, B(xB4, B5)=1
Japanese (Miyazaki)0.090100 [15] B4b1a1=3, B4(xB4a, B4b1)=2, B4a=2, B5b=2
Khakassian (Khakassia)0.08857 [6] B4=5
Han (Xinjiang)0.08547[ citation needed ]B5a=2, B4b1=1, B5b=1
Kyrgyz (Sary-Tash, Kyrgyzstan)0.08547[ citation needed ]B5a=2, B4(xB4a, B4b1, B4c)=1, B4a=1
Mongolian (Ulan Bator)0.08547 [10] B4(xB4a, B4b, B4c)=2, B4a=1, B5b=1
Tuvan 0.08495 [14] B4a1c2=4, B5=2, B4(xB4a1c2, B4b1a, B4d1a)=1, B4b1a=1
Tibetan (Nyingchi, Tibet)0.08324[ citation needed ]B=2
Barghut (Hulunbuir)0.081149 [16] B4c1a2(xB4c1a2a)=4, B4f1=2, B5b(xB5b2)=2, B4c1a2a=1, B4b1a3a1a=1, B4d1=1, B5b2=1
Chamorro (85 Guam, 14 Saipan, & 6 Rota)0.076105[ citation needed ]B4a1a1a=7, B4b1=1
Manchurian 0.07540 [10] B4(xB4a, B4b, B4c)=2, B4a=1
Uzbek (Uzbekistan/Kyrgyzstan)0.07540[ citation needed ]B=3
Uyghur (Penjim, Panfilov District, Almaty Province, Kazakhstan)0.07355[ citation needed ]B5a=3, B4(xB4a, B4b1, B4c)=1
Buryat 0.071126[ citation needed ]B4(xB4a, B4b)=5, B5b=2, B(xB4, B5a, B5b)=1, B4b=1
Khamnigan (Buryatia)0.07199 [6] B4=5, B5=2
Dungan (Uzbekistan/Kyrgyzstan)0.06316[ citation needed ]B=1
Persian (eastern Iran)0.06182 [6] B4=4, B5=1
Nogai (Dagestan)0.06133[ citation needed ]B=2
Tibetan (Nagchu, Tibet)0.05735[ citation needed ]B=2
Kazakh (Kegen Valley, Almaty Province, Kazakhstan)0.05555[ citation needed ]B4(xB4a, B4b1, B4c)=1, B4a=1, B5b=1
Lisu (Gongshan, Yunnan)0.05437[ citation needed ]B4(xB4a, B4b1)=1, B5(xB5a, B5b)=1
Tharu (Chitwan, Nepal)0.053133[ citation needed ]B5a=7
Uzbek (Xinjiang)0.05258[ citation needed ]B4b1=2, B4c=1
Bukharan Arab (Uzbekistan/Kyrgyzstan)0.05020[ citation needed ]B=1
Filipino (Palawan)0.05020[ citation needed ]B4a1a=1
Tajik (Uzbekistan/Kyrgyzstan)0.05020[ citation needed ]B=1
Turkmen (Uzbekistan/Kyrgyzstan)0.05020[ citation needed ]B=1
Shor (Kemerovo Oblast)0.04982 [6] B4=4
Va (Simao, Yunnan)0.04522[ citation needed ]B4(xB4a)=1
Altai Kizhi 0.04490 [6] B4=3, B5=1
Bai (Dali, Yunnan)0.04468[ citation needed ]B5a=2, B4b1=1
Tofalar 0.04346 [14] B4a1c2=2
Kyrgyz (Bakay-Ata, Kyrgyzstan)0.04248[ citation needed ]B4(xB4a, B4b1, B4c)=1, B5b=1
Tubalar 0.04272 [14] B4b1a=3
Evenk (Krasnoyarsk Krai)0.04173 [6] B4=3
Kazakh (Kosh-Agach, Altai Republic)0.04198 [16] B5b2=3, B4c1b=1
Buryat (Kushun, Nizhneudinsky, Irkutsk Oblast)0.04025 [14] B4d1a=1
Kazakh (Xinjiang)0.03853[ citation needed ]B4a=2
Teleut (Kemerovo Oblast)0.03853 [6] B4=2
Kalmyk (Kalmykia)0.036110 [6] B4=3, B5=1
Jetisu Kazakhstan 0.035200 [19] B4=4, B5=3
Tibetan (Chamdo, Tibet)0.03429[ citation needed ]B4a=1
Buryat (Buryatia)0.034295 [6] B4=9, B5=1
Lahu (Lancang, Yunnan)0.02935[ citation needed ]B4(xB4a, B4b1)=1
Pumi (Ninglang, Yunnan)0.02836[ citation needed ]B4(xB4a, B4b1)=1
Va (Ximeng & Gengma, Yunnan)0.02836[ citation needed ]B5a=1
Yakut (Yakutia)0.02836 [6] B4=1
Tharu (Morang, Nepal)0.02540[ citation needed ]B5a=1
Tibetan (Deqin, Yunnan)0.02540[ citation needed ]B4(xB4a, B4b1)=1
Oroqen (Oroqen Autonomous Banner)0.02344[ citation needed ]B4(xB4a, B4b)=1
Uyghur (Xinjiang)0.02147[ citation needed ]B(xB4, B5, B6)=1
Oirat Mongol (Xinjiang)0.02049[ citation needed ]B4b1=1
Ainu 0.02051[ citation needed ]B4f1=1
Tuvinian (Tuva)0.019105 [6] B4=2
Tibetan (Qinghai)0.01856[ citation needed ]B4a=1
Nogai (Nogaysky, Dagestan & Adyge-Khabalsky, Karachay-Cherkessia)0.015206[ citation needed ]B=3
Tibetan (Shannan, Tibet)0.01474[ citation needed ]B4a=1
Yakut 0.012423[ citation needed ]B4(xB4b1)=3
B4b1=1
B5b2=1
Bashkir 0.009221[ citation needed ]B=2
Chukchi (Anadyr)0.00015 [6] –
Uyghur (Uzbekistan/Kyrgyzstan)0.00016[ citation needed ]–
Crimean Tatar (Uzbekistan/Kyrgyzstan)0.00020[ citation needed ]–
Iranian (Uzbekistan/Kyrgyzstan)0.00020[ citation needed ]–
Karakalpak (Uzbekistan/Kyrgyzstan)0.00020[ citation needed ]–
Kazakh (Uzbekistan/Kyrgyzstan)0.00020[ citation needed ]–
Kyrgyz (Uzbekistan/Kyrgyzstan)0.00020[ citation needed ]–
Hindu (Chitwan, Nepal)0.00024[ citation needed ]–
Nganasan 0.00024 [14] –
Tibetan (Diqing, Yunnan)0.00024[ citation needed ]–
Kurd (northwestern Iran)0.00025 [6] –
Andhra Pradesh (tribal)0.00029[ citation needed ]–
Tibetan (Shigatse, Tibet)0.00029[ citation needed ]–
Batak (Palawan)0.00031[ citation needed ]–
Ket 0.00038 [14] –
Tajik (Tajikistan)0.00044 [6] –
Tibetan (Lhasa, Tibet)0.00044[ citation needed ]–
Evenk (Buryatia)0.00045 [6] –
Udege (Gvasiugi, Imeni Lazo, Khabarovsk Krai)0.00046 [14] –
Itelmen 0.00047 [14] –
Chuvash 0.00055[ citation needed ]–
Nivkh (northern Sakhalin)0.00056 [14] –
Komi-Zyryans 0.00062[ citation needed ]–
Mansi 0.00063[ citation needed ]–
Chukchi 0.00066 [14] –
Evenk (53 Stony Tunguska basin & 18 Tuguro-Chumikan)0.00071 [14] –
Tatar (Aznakayevo)0.00071[ citation needed ]–
Komi-Permyaks 0.00074[ citation needed ]–
Siberian Eskimo 0.00079 [14] –
Ulchi 0.00087 [14] –
Mansi 0.00098 [14] –
Udmurt 0.000101[ citation needed ]–
Mordvinian 0.000102[ citation needed ]–
Khanty 0.000106[ citation needed ]–
Yakut 0.000117[ citation needed ]–
Tatar (Buinsk)0.000126[ citation needed ]–
Mari 0.000136[ citation needed ]–
Koryak 0.000155 [14] –
Tatar 0.000228[ citation needed ]–

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.

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. S2CID   27566749.
  2. ↑ Yao, Yong-Gang; et al. (March 2002). "Phylogeographic Differentiation of Mitochondrial DNA in Han Chinese". Am J Hum Genet. 70 (3): 635–651. doi:10.1086/338999. PMC   384943 . PMID   11836649.
  3. ↑ "Downloadable genotypes of present-day and ancient DNA data (compiled from published papers)". reich.hms.harvard.edu. David Reich Lab. Archived from the original on 2019-11-02. Retrieved 2019-09-11.
  4. ↑ Haplogroup B.
  5. ↑ Fagundes, Nelson J.R.; Kanitz, Ricardo; Eckert, Roberta; Valls, Ana C.S.; Bogo, Mauricio R.; Salzano, Francisco M.; Smith, David Glenn; Silva, Wilson A.; Zago, Marco A.; Ribeiro-dos-Santos, Andrea K.; Santos, Sidney E.B.; Petzl-Erler, Maria Luiza; Bonatto, Sandro L. (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.
  6. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Derenko, Miroslava; Malyarchuk, Boris; Grzybowski, Tomasz; et al. (2007). "Phylogeographic Analysis of Mitochondrial DNA in Northern Asian Populations". Am. J. Hum. Genet. 81 (5): 1025–1041. doi:10.1086/522933. PMC   2265662 . PMID   17924343.
  7. ↑ Al-Zahery, Nadia; Pala, Maria; Battaglia, Vincenza; et al. (2011). "In search of the genetic footprints of Sumerians: A survey of Y-chromosome and mtDNA variation in the Marsh Arabs of Iraq". BMC Evolutionary Biology. 2011 (11): 288. Bibcode:2011BMCEE..11..288A. doi: 10.1186/1471-2148-11-288 . PMC   3215667 . PMID   21970613.
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