Haplogroup U (mtDNA)

Haplogroup U
Possible time of origin 46,500 ± 3,300 years ago[1]
Possible place of origin Western Asia[2]
Ancestor R
Descendants U1, U5, U6, U2'3'4'7'8'9
Defining mutations 11467, 12308, 12372[3]

Haplogroup U is a human mitochondrial DNA haplogroup (mtDNA). Arising from haplogroup R in the beginning Upper Paleolithic, its various subclades (labelled U1–U9, diverging over the course of the Upper Paleolithic) are found widely distributed across Northern and Eastern Europe, Central, Western and South Asia as well as North Africa and the Horn of Africa.

The most common subclade of haplogroup U8b, arising between about 30,000 and 22,000 years ago, is given the separate designation of haplogroup K.

Origins

Haplogroup U descends from the haplogroup R mtDNA branch of the phylogenetic tree. The defining mutations (A11467G, A12308G, G12372A) are estimated to have arisen between about 43,000 and 50,000 years ago, in the beginning Upper Paleolithic (95% CI: 46,530 ± 3,290 according to Behar et al., 2012)

Ancient DNA classified as belonging to the U* mitochondrial haplogroup has been recovered from human remains found in Western Siberia, dated to c. 45,000 years ago..[4]

Haplogroup U has also been found among ancient Egyptian mummies excavated at the Abusir el-Meleq archaeological site in Middle Egypt, dated to the 1st millennium BC.[5]

Distribution

Haplogroup U is found in 15% of Indian caste and 8% of Indian tribal populations.[6] Haplogroup U is found in approximately 11% of native Europeans and is held as the oldest maternal haplogroup found in that region.[6][7][8] In a 2013 study, all but one of the ancient modern human sequences from Europe belonged to maternal haplogroup U, thus confirming previous findings that haplogroup U was the dominant type of Mitochondrial DNA (mtDNA) in Europe before the spread of agriculture into Europe and the presence and the spread of the Indo-Europeans in Western Europe.[9][10]

Haplogroup U has various subclades numbered U1 to U9. Haplogroup K is a subclade of U8.[11] The old age has led to a wide distribution of the descendant subgroups across Western Eurasia, North Africa, and South Asia. Some subclades of haplogroup U have a more specific geographic range.

Subclades

Subclades are labelled U1–U9; Haplogroup K is a subclade of U8.

Van Oven and Kayser (2009) proposed subclades "U2'3'4'7'8" and "U4'9".[3] Behar et al. (2012) amended this by grouping "U4'9" as subordinate to "U2'3'4'7'8" for a new intermediate subclade "U2'3'4'7'8'9".

Haplogroup U1

Haplogroup U1 estimated to have arisen between 26,000 and 37,000 years ago. It is found at very low frequency throughout Europe. It is more often observed in eastern Europe, Anatolia and the Near East. It is also found at low frequencies in India. U1 is found in the Svanetia region of Georgia at 4.2%. Subclade U1a is found from India to Europe, but is extremely rare among the northern and Atlantic fringes of Europe including the British Isles and Scandinavia. Several examples in Tuscany have been noted. In India, U1a has been found in the Kerala region. U1b has a similar spread but is rarer than U1a. Some examples of U1b have been found among Jewish diaspora. Subclades U1a and U1b appear in equal frequency in eastern Europe.[12]

The U1 subclades are: U1a (with deep-subclades U1a1, U1a1a, U1a1a1, U1a1b)[13] and U1b.[13]

DNA analysis of excavated remains now located at ruins of the Church of St. Augustine in Goa, India revealed the unique mtDNA subclade U1b, which is absent in India, but present in Georgia and surrounding regions.[14] Since the genetic analysis corroborates archaeological and literary evidence, it is believed that the excavated remains belong to Ketevan the Martyr, queen of Georgia.[14]

Haplogroup U1 has been observed among specimens at the mainland cemetery in Kulubnarti, Sudan, which date from the Early Christian period (AD 550-800).[15]

The rare U1 clade is also found among Algerians in Oran (0.83%-1.08%) and the Reguibat tribe of the Sahrawi (0.93%).[16]

Haplogroup U5

The age of U5 is estimated at between 25,000 and 35,000 years old.[17] Approximately 11% of total Europeans and 10% of European-Americans are in haplogroup U5.

U5 has been found in human remains dating from the Mesolithic in England, Germany, Lithuania, Poland, Portugal, Russia,[18] Sweden,[19] France[20] and Spain.[21] Neolithic skeletons (~7,000 years old) that were excavated from the Avellaner cave in Catalonia, northeastern Spain included a specimen, which also carried haplogroup U5.[22]

Haplogroup U5 and its subclades U5a and U5b today form the highest population concentrations in the far north, among Sami, Finns, and Estonians. However, it is spread widely at lower levels throughout Europe. This distribution, and the age of the haplogroup, indicate individuals belonging to this clade were part of the initial expansion tracking the retreat of ice sheets from Europe around 10,000 years ago.

Additionally, haplogroup U5 is found in small frequencies and at much lower diversity in the Near East and parts of northern Africa (areas with sizable U6 concentrations), suggesting back-migration of people from Europe toward the south.[23]

Mitochondrial haplogroup U5a has also been associated with HIV infected individuals displaying accelerated progression to AIDS and death.[24]

Haplogroup U6

Projected frequencies for haplogroup U6 (top left) and several subclades.

Haplogroup U6 is dated to between 31,000 and 43,000 years ago by Behar et al. (2012). This is consistent with the discovery of basal U6* in a Romanian specimen of ancient DNA (Peștera Muierilor) dated to 35,000 years ago.[41] Hervella et al. (2016) take this find as evidence for Paleolithic back-migration of Homo sapiens from Eurasia into Africa. The discovery of basal U6* in ancient DNA contributed to setting back the estimated age of U6 to around 46,000 years ago. [42]

Haplogroup U6 is common (with a prevalence of around 10%)[23] in Northwest Africa (with a maximum of 29% in an Algerian Mozabites[43]) and the Canary Islands (18% on average with a peak frequency of 50.1% in La Gomera). It is also found in the Iberian peninsula, where it has the highest diversity (10 out of 19 sublineages are only found in this region and not in Africa),[44] Northeast Africa and occasionally in other locations. U6 is also found at low frequencies in the Chad Basin, including the rare Canarian branch. This suggests that the ancient U6 clade bearers may have inhabited or passed through the Chad Basin on their way westward toward the Canary Islands.[45]

U6 is thought to have entered North Africa from the Near East around 30,000 years ago. It has been found among Iberomaurusian specimens dating from the Epipaleolithic at the Taforalt prehistoric site.[46] In spite of the highest diversity of Iberian U6, Maca-Meyer argues for a Near East origin of this clade based on the highest diversity of subclade U6a in that region,[44] where it would have arrived from West Asia, with the Iberian incidence primarily representing migration from the Maghreb and not persistence of a European root population.

According to Hernández et al. 2015 "the estimated entrance of the North African U6 lineages into Iberia at 10 ky correlates well with other L African clades, indicating that U6 and some L lineages moved together from Africa to Iberia in the Early Holocene."[47]

U6 has four main subclades:[44][46]

Subgroup U6a reflects the first African expansion from the Maghrib returning to the east. Derivative clade U6a1 signals a posterior movement from East Africa back to the Maghrib and the Near East. This migration coincides with the probable Afroasiatic linguistic expansion. U6b and U6c clades, restricted to West Africa, had more localized expansions. U6b probably reached the Iberian Peninsula during the Capsian diffusion in North Africa. Two autochthonous derivatives of these clades (U6b1 and U6c1) indicate the arrival of North African settlers to the Canarian Archipelago in prehistoric times, most probably due to the Saharan desiccation. The absence of these Canarian lineages nowadays in Africa suggests important demographic movements in the western area of this Continent.
— Maca-Meyer 2003

U6a, U6b and U6d share a common basal mutation (16219) that is not present in U6c, whereas U6c has 11 unique mutations. U6b and U6d share a mutation (16311) not shared by U6a, which has three unique mutations.

U2'3'4'7'8'9

Subclades U2, U3, U4, U7, U8 and U9 are now thought to be monophyletic, its common ancestor "U2'3'4'7'8'9" defined by mutation A1811G, arising between about 42,000 and 48,000 years ago (Behar et al., 2012). Within U2'3'4'7'8'9, U4 and U9 may be monophyletic, as "U4'9" (mutations T195C!, G499A, T5999C) arising between 31,000 and 43,000 years ago (Behar et al., 2012).

Haplogroup U2

Haplogroup U2 is most common in South Asia[49] but is also found in low frequency in Central and West Asia, as well as in Europe as U2e (the European variety of U2 is named U2e).[50] The overall frequency of U2 in South Asia is largely accounted for by the group U2i in India whereas haplogroup U2e, common in Europe, is entirely absent; given that these lineages diverged approximately 50,000-years-ago, these data have been interpreted as indicating very low maternal-line gene-flow between South Asia and Europe throughout this period.[49] Approximately one half of the U mtDNAs in India belong to the Indian-specific branches of haplogroup U2 (U2i: U2a, U2b and U2c).[49] While U2 is typically found in India, it is also present in the Nogais, descendants of various Mongolic and Turkic tribes, who formed the Nogai Horde.[51] Both U2 and U4 are found in the Ket and Nganasan peoples, the indigenous inhabitants of the Yenisei River basin and the Taymyr Peninsula.[52]

The U2 subclades are: U2a,[53] U2b,[54] U2c,[55] U2d,[56] and U2e.[57] With the India-specific subclades U2a, U2b, and U2c collectively referred to as U2i, the Eurasian haplogroup U2d appears to be a sister clade with the Indian haplogroup U2c,[58] while U2e is considered a European-specific subclade but also found in South India.[50][59]

Haplogroup U2 has been found in the remains of a 30,000-year-old hunter-gatherer from the Kostyonki, Voronezh Oblast in Central-South European Russia.,[60] in 4800 to 4000-year-old human remains from a Beaker culture site of the Late Neolithic in Kromsdorf Germany,[61] and in 2,000-year-old human remains from Bøgebjerggård in Southern Denmark. However, haplogroup U2 is rare in present-day Scandinavians.[62] The remains of a 2,000-year-old West Eurasian male of haplogroup U2e1 was found in the Xiongnu Cemetery of Northeast Mongolia.[63]

Haplogroup U3

Haplogroup U3 falls into two subclades:: U3a[64] and U3b.[64]

Coalescence age for U3a is estimated as 18,000 to 26,000-years-ago while the coalescence age for U3b is estimated as 18,000 to 24,000-years-ago. U3a is found in Europe, the Near East, the Caucasus and North Africa. The almost-entirely European distributed subclade, U3a1, dated at 4000 to 7000-years-ago, suggests a relatively recent (late Holocene or later) expansion of these lineages in Europe. There is a minor U3c subclade (derived from U3a), represented by a single Azeri mtDNA from the Caucasus. U3b is widespread across the Middle East and the Caucasus, and it is found especially in Iran, Iraq and Yemen, with a minor European subclade, U3b1b, dated at 2000 to 3000-years-ago.[65] Haplogroup U3 is defined by the HVR1 transition A16343G. It is found at low levels throughout Europe (about 1% of the population), the Near East (about 2.5% of the population), and Central Asia (about 1% of the population). U3 is present in the Svan population from the Svaneti region (about 4.2% of the population) and among Lithuanian Romani, Polish Romani, and Spanish Romani populations (36-56%)[66][67][25] consistent with a common migration route from India then out-of-the Balkans for the Lithuanian, Polish, and Spanish Roma.[68]

The U3 clade is also found among Mozabite Berbers (10.59%),[16] as well as Egyptians in the El-Hayez (2.9%)[69] and Gurna oases (2.9%),[70] and Algerians in Oran (1.08%-1.25%).[16] The rare U3a subclade occurs among the Tuareg inhabiting Niger (3.23%).[71]

Haplogroup U3 has been found in some of the 6400-year-old remains (U3a) discovered in the caves at Wadi El‐Makkukh near Jericho associated with the Chalcolithic period.[72] Haplogroup U3 was already present in the West Eurasian gene pool around 6,000-years-ago and probably also its subclade U3a as well.[72]

Haplogroup U4

Haplogroup U4 has its origin between 21,000 and 14,000 years ago. Its distribution is associated with the population bottleneck due to the Last Glacial Maximum.[73]

U4 has been found in ancient DNA,[74] and it is relatively rare in modern populations,[75] although it is found in substantial ratios in certain indigenous populations of Northern Asia and Northern Europe, being associated with the remnants of ancient European hunting-gatherers preserved in the indigenous populations of Siberia.[76][77][78] U4 is found in the Nganasan people of the Taymyr Peninsula,[52][79] in the Mansi (16.3%) an endangered people,[78] and in the Ket people (28.9%) of the Yenisei River.[78] It is found in Europe with highest concentrations in Scandinavia and the Baltic states.[80] and is found in the Sami population of the Scandinavian peninsula (although, U5b has a higher representation).[81][82] U4 is also preserved in the Kalash people (current population size 3,700)[83] a unique tribe among the Indo-Aryan peoples of Pakistan where U4 (subclade U4a1[84]) attains its highest frequency of 34%.[85][86][87]

The U4 subclades are: U4a,[88] U4b,[89] U4c,[90] and U4d.[91]

Haplogroup U4 is associated with ancient European hunter-gatherers and has been found in 7,200 to 6,000-year-old remains of the Pitted Ware culture in Gotland Sweden and in 4,400 to 3,800-year-old remains from the Damsbo site of the Danish Beaker culture.[92][19][93] Remains identified as subclade U4a2 are associated with the Corded Ware culture, which flourished 5200 to 4300 years ago in Eastern and Central Europe and encompassed most of continental northern Europe from the Volga River in the east to the Rhine in the west.[94] Mitochondrial DNA recovered from 3,500 to 3,300-year-old remains at the Bredtoftegård site in Denmark associated with the Nordic Bronze Age include haplogroup U4 with 16179T in its HVR1 indicative of subclade U4c1.[93][95][96][97]

Haplogroup U7

Haplogroup U7 is considered a West Eurasian-specific mtDNA haplogroup, believed to have originated in the Black Sea area approximately 30,000-years-ago.[98][99][100] In modern populations, U7 occurs at low frequency in the Caucasus,[100] the western Siberian tribes,[101] West Asia (about 4% in the Near East, while peaking with 10% in Iranians),[98] South Asia (about 12% in Gujarat, the westernmost state of India, while for the whole of India its frequency stays around 2%, and 5% in Pakistan),[98] and the Vedda people of Sri Lanka where it reaches it highest frequency of 13.33% (subclade U7a).[102] One third of the West Eurasian-specific mtDNAs found in India are in haplogroups U7, R2 and W. It is speculated that large-scale immigration carried these mitochondrial haplogroups into India.[98]

The U7 subclades are: U7a (with deep-subclades U7a1, U7a2, U7a2a, U7a2b)[103] and U7b.[103]

Genetic analysis of individuals associated with the Late Hallstatt culture from Baden-Württemberg Germany considered to be examples of Iron Age "princely burials" included haplogroup U7.[104] Haplogroup U7 was reported to have been found in 1200-year-old human remains (dating to around 834), in a woman believed to be from a royal clan who was buried with the Viking Oseberg Ship in Norway.[105] Haplogroup U7 was found in 1000-year-old human remains (dating to around AD 1000-1250) in a Christian cemetery is Kongemarken Denmark. However, U7 is rare among present-day ethnic Scandinavians.[101]

The U7a subclade is especially common among Saudis, constituting around 30% of maternal lineages in the Eastern Province.[106]

The subclade U7a4 point to an origin in the Near East. It peaks among the modern inhabitants of Azerbaijan (26%) and Iranian Azerbaijanis (16-22%), while occurring in the rest of Iran at frequencies from 2-16%, with moderate frequencies of 14-17% in the western parts of Iraq and 1-19% in the Arabian Peninsula. In lower frequencies it is also found in Turkmenistan (2-10%), western parts of Kazakhstan (2-10%), Syria (3-7%), Uzbekistan (2-6%), Central Anatolia (2-4%), South Punjab (1-2%), and Afghanistan (0-2%). U7a4 was also observed in one instance in Tuscany and two other in European Russia.

Haplogroup U8

Haplogroup U9

Haplogroup U9 is a rare clade in mtDNA phylogeny, characterized only recently in a few populations of Pakistan (Quintana-Murci et al. 2004). Its presence in Ethiopia and Yemen, together with some Indian-specific M lineages in the Yemeni sample, points to gene flow along the coast of the Arabian Sea. Haplogroups U9 and U4 share two common mutations at the root of their phylogeny. It is interesting that, in Pakistan, U9 occurs frequently only among the so-called Makrani population. In this particular population, lineages specific to parts of Eastern Africa occur as frequently as 39%, which suggests that U9 lineages in Pakistan may have an origin from this area (Quintana-Murci et al. 2004). Regardless of which coast of the Arabian Sea may have been the origin of U9, its Ethiopian–southern Arabian–Indus Basin distribution hints that the subclade's diversification from U4 may have occurred in regions far away from the current area of the highest diversity and frequency of haplogroup U4—East Europe and western Siberia.[107]

See also

Wikimedia Commons has media related to Haplogroup U (mtDNA).

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. ↑ Behar et al. (2012), note the revised estimate of 42–58 kya based on ancient DNA by Hervella et al. (2016).
  2. ↑ Hervella et al. (2016) "Individuals carrying haplogroup U possibly spread westward from Western Asia around 39–52 ky, reaching Europe as signaled by haplogroup U5, and North Africa signaled by haplogroup U6"
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  4. ↑ "Carriers of mitochondrial DNA macrohaplogroup R colonized Eurasia and Australasia from a southeast Asia core area". BMC Evolutionary Biology. 17. doi:10.1186/s12862-017-0964-5.. "Carriers of human mitochondrial DNA macrohaplogroup M colonized India from southeastern Asia". BMC Evolutionary Biology. 16. doi:10.1186/s12862-016-0816-8.
  5. ↑ Schuenemann, Verena J.; et al. (2017). "Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods". Nature Communications. 8: 15694. PMID 28556824. doi:10.1038/ncomms15694.
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  12. ↑ , FamilyTreeDNA - mtDNA haplogroup U1 project
  13. 1 2 , Phylogeny of mt-hg U3, Ron Scott 2010
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  15. ↑ Sirak, Kendra; Frenandes, Daniel; Novak, Mario; Van Gerven, Dennis; Pinhasi, Ron (2016). Abstract Book of the IUAES Inter-Congress 2016 - A community divided? Revealing the community genome(s) of Medieval Kulubnarti using next- generation sequencing. IUAES.
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  18. ↑ Bramanti B, Thomas MG, Haak W, et al. (October 2009). "Genetic discontinuity between local hunter-gatherers and central Europe's first farmers". Science. 326 (5949): 137–40. PMID 19729620. doi:10.1126/science.1176869.
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  23. 1 2 The Genographic Project at National Geographic
  24. 1 2 Hendrickson SL, Hutcheson HB, Ruiz-Pesini E, et al. (November 2008). "Mitochondrial DNA haplogroups influence AIDS progression". AIDS. 22 (18): 2429–39. PMC 2699618 Freely accessible. PMID 19005266. doi:10.1097/QAD.0b013e32831940bb.
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  26. ↑ Age: 1,398.1 ± 1,872.3; CI=95% (Behar et al., 2012) haplogroup.org
  27. ↑ 22,794.0 ± 3,590.3; CI=95% (Behar et al., 2012) haplogroup.org
  28. ↑ C. Coudray; A. Olivieri; A. Achilli; M. Pala; M. Melhaoui; M. Cherkaoui; F. El-Chennawi; M. Kossmann; A. Torroni; J. M. Dugoujon (August 2009). "The Complex and Diversified Mitochondrial Gene Pool of Berber Populations". Annals of Human Genetics. 73 (2): 196–214. PMID 19053990. doi:10.1111/j.1469-1809.2008.00493.x. Retrieved 7 July 2015.
  29. ↑ Age: 15,529.7 ± 4,889.9; CI=95% (Behar et al., 2012) haplogroup.org
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  33. ↑ 20,040.2 ± 3,208.6; CI=95% (Behar et al., 2012) haplogroup.org
  34. ↑ 14,938.0 ± 3,656.4; CI=95% (Behar et al., 2012) haplogroup.org
  35. 1 2 3 Boris Malyarchuk et al., "The Peopling of Europe from the Mitochondrial Haplogroup U5 Perspective", PLoS One. 2010; 5(4): e10285. 10.1371/journal.pone.0010285. "subcluster U5b2a is characterized by a predominantly central European distribution, since a large number of U5b samples from Poland, Slovakia and the Czech Republic fall into this subcluster. For instance, subcluster U5b2a2 is frequent in central Europe (with the highest frequency of its subcluster U5b2a2a1 in Poles) and dated as arising between 12–18 kya [...] It is also remarkable that within U5b2a1a, a Mediterranean branch precedes subcluster U5b2a1a1, which is characteristic of central and eastern European populations (Figure 1). Another U5b subcluster, U5b3, has its most likely homeland in the Italian Peninsula, from where it expanded in the Holocene along the Mediterranean coasts [11]. Hence, in general one may conclude that an initial diversification of U5b occurred in southern and central Europe, followed by the spread of a particular U5b subclusters into eastern Europe."
  36. ↑ Age: 14,676.9 ± 2,716.6; CI=95% (Behar et al., 2012) haplogroup.org
  37. ↑ Age: 12,651.1 ± 5,668.2; CI=95% (Behar et al., 2012) haplogroup.org
  38. ↑ 4,015.9 ± 3,898.7; CI=95% (Behar et al., 2012) haplogroup.org
  39. ↑ The Young Man of Byrsa specimen dates from the late 6th century BCE, and his lineage is believed to represent early gene flow from Iberia to the Maghreb. Matisoo-Smith EA, Gosling AL, Boocock J, Kardailsky O, Kurumilian Y, Roudesli-Chebbi S, et alMatisoo-Smith EA; Gosling AL; Boocock J; Kardailsky O; Kurumilian Y; Roudesli-Chebbi S; et al. (May 25, 2016). "A European Mitochondrial Haplotype Identified in Ancient Phoenician Remains from Carthage, North Africa" (PDF). PLoS ONE. 11 (5): e0155046. PMC 4880306 Freely accessible. PMID 27224451. doi:10.1371/journal.pone.0155046. Retrieved 26 May 2016.
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  49. 1 2 3 , Most of the extant mtDNA boundaries in South and Southwest Asia were likely shaped during the initial settlement of Eurasia by anatomically modern humans, Mait Metspalu, et al. 2004
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  54. ↑ , Phylogeny of mt-hg U2b, Ron Scott 2010
  55. ↑ , Phylogeny of mt-hg U2c, Ron Scott 2009
  56. ↑ , Phylogeny of mt-hg U2d, Ron Scott 2009
  57. ↑ , Phylogeny of mt-hg U2e, Ron Scott 2010
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  64. 1 2 , Phylogeny of mt-hg U3, Ron Scott 2010
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  74. ↑ , TNeolithic pots and potters in Europe: the end of ‘demic diffusion’ migratory mode, Mihael Budja 2013
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  77. ↑ , Ancient DNA from European Early Neolithic Farmers Reveals Their Near Eastern Affinities, Haak et al. 2010
  78. 1 2 3 , Traces of Early Eurasians in the Mansi of Northwest Siberia Revealed by Mitochondrial DNA, Derbeneva et al. 2002
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  80. ↑ , Maternal Ancestry: The clan of Ulrike, Oxford Ancestors, 2013
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  82. ↑ , Oxford Ancestors Ltd. - Maternal Ancestry
  83. ↑ , Pakistan Today: Safeguarding Kalash heritage, Bashir, 2012
  84. ↑ , U4a1 Genbank Sequences, Ian Logan 2014
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  87. ↑ , U4 Haplogroup: A blog dedicated to the U4 Haplogroup - The Kalash,
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  89. ↑ , Phylogeny of mt-hg U4b, Ron Scott 2011
  90. ↑ , Phylogeny of mt-hg U4c, Ron Scott 2011
  91. ↑ , Phylogeny of mt-hg U4d, Ron Scott 2010
  92. ↑ , Eupedia: Prehistoric European DNA - mtDNA & Y-DNA Haplogroup Frequencies by Period
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  96. ↑ , Genetic Diversity among Ancient Nordic Populations: Table 1. Nucleotide substitutions and mtDNA haplogroups assigned for individuals from the Neolithic site Damsbo (4,200 YBP) and the Early Bronze Age site Bredtoftegård (3,300–3,500 YBP), Melchior et al 2010
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  98. 1 2 3 4 , Most of the extant mtDNA boundaries in South and Southwest Asia were likely shaped during the initial settlement of Eurasia by anatomically modern humans. Metspalu et al., 2004.
  99. ↑ , The Oseberg women and the Gokstad man retain their genetic secrets, Kulturhistorisk Museum - University of Oslo
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