→Origins: Cleaned up origins, removed information redundant with R1a1a page. |
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As of November 2009 R1a is defined by the following mutations: L62?, L63?, M420, M449, M511, M513. |
As of November 2009 R1a is defined by the following mutations: L62?, L63?, M420, M449, M511, M513. |
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The exact point of origin of R1a's dispersal pattern found today remains the subject of discussion. It presumably originated somewhere in the [[Eurasia]]n landmass, where it is most commonly found today. There |
The exact point of origin of R1a's dispersal pattern found today remains the subject of discussion. It presumably originated somewhere in the [[Eurasia]]n landmass, where it is most commonly found today. There is a broad discussion of the possible origin of the largest subclade within R1a*/R1a1* tiers, [[Haplogroup R1a1a (Y-DNA)|R1a1a]]. Establishing the origin of this haplogroup maybe considerably easier than establishing the place of origin of R1a* since R1a* is relatively scarce. |
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Because R1a* and R1a1* are spread at low frequencies across a broad area, it is difficult to pinpoint sites of diversity, while many studies point toward R1a1a diversity, it should be remembered that R1a1a arose from a single R1a1 Y chromosome and could have done so anywhere that R1a1 had spread at the time R1a1a evolved. {{Harvcoltxt|Sharma et al.|2009}} indicated a site of basal R1a diversity around Bhopal, India with 13 individuals bearing the R1a* mutation, however unlike other study subjects, these R1a were not included as part of the more comprehensive study of Underhill et al. (2009). {{Harvcoltxt|Mirabal et al.|2009}}, and {{Harvcoltxt|Underhill et al.|2009}}, concluded that there are two separate "poles of the expansion" with similar ages, with South Asian R1a older than European R1a. Of 117 R1a examined from India, none where R1a* or R1a1* (all were R1a1a) contradicting Sharma et al. 2009. |
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In 2009, two large studies of available data, {{Harvcoltxt|Mirabal et al.|2009}}, and {{Harvcoltxt|Underhill et al.|2009}}, concluded that there are two separate "poles of the expansion" with similar ages, with South Asian R1a older than European R1a. While neither group of authors took a decisive position, both articles felt the data consistent with South Asian origins. {{Harvcoltxt|Mirabal et al.|2009}} felt the data to also be consistent with Central Asian origins, while {{Harvcoltxt|Underhill et al.|2009}} took to be also consistent with Western Asian origins. In any case, the publication of these major articles, both with large panels of co-authors, makes the Asian origins the leading theory as of late 2009. |
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{{further|[[Haplogroup R1a1a (Y-DNA)]]}} |
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===Central Asian Origin Theories=== |
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{{Harvcoltxt|Cordaux et al.|2004}} argued, citing data from {{Harvcoltxt|Wells et al.|2001}}, {{Harvcoltxt|Semino et al.|2000}}, and {{Harvcoltxt|Quintana-Murci et al.|2001}} that... |
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{{cquote|Given the high frequency of R-M17 in [[central Asia]] (typically 20%–40% [9]), its rarity in [[west Asia]] [9, 13] and its absence in east Asia [14], Indian R-M17 Y chromosomes most probably have a central Asian origin [8, 9].|{{Harvcoltxt|Cordaux et al.|2004}}}} |
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This position is also considered likely by {{Harvcoltxt|Mirabal et al.|2009}} after their larger analysis of recent data. |
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===Eastern European Origin Theories=== |
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Suggestions have been made which associate the distribution of R1a with several proposed movements of people in history and prehistory in [[Eastern Europe]] :- |
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*'''End of the [[Ice Age]].''' The spread from a Ukrainian refugium during the [[Late Glacial Maximum]] |
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*'''[[Bronze Age]].''' The spread of [[Indoeuropean languages]] and/or [[Indo-Aryan languages]] and/or [[Indo-Iranian languages]] in the [[Bronze Age]] (also associated with the use of [[horses]], and "[[Kurgan]]" burials). |
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*'''Historical Era.''' The spread of [[Slavic languages]] and migrations in [[late Classical]] times. |
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These three proposals involve very different time periods, but they are not mutually exclusive given that R1a lineages may have been taken part in many different human movements over time in the same geographical region.<ref>{{Harvcoltxt|Semino et al.|2000}} proposed quite early that there may have been two expansions, suggesting that the spread of R1a from [[Ukrainian LGM refuge|a point of origin in Ukraine]] following the [[Last Glacial Maximum]] may have been magnified by the expansion of males from the [[Kurgan culture]]. In a study of the Balkans, {{Harvcoltxt|Pericic et al.|2005}} saw evidence for "at least three major episodes of gene flow" adding "possibly massive Slavic migration from A.D. 5th to 7th centuries" as a third.</ref> |
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====End of the Ice Age==== |
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Age estimates of this depth come from papers using the methodology described by {{Harvcoltxt|Zhivotovsky et al.|2004}}, the latest such example being {{Harvcoltxt|Mirabal et al.|2009}} and {{Harvcoltxt|Underhill et al.|2009}}. Researchers using this estimation method therefore believe any Bronze Age or more recent dispersals affecting modern R1a diversity must be specific to certain sub-clades, such as R-M458. |
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====Bronze Age (Indo Europeans, Indo-Aryans, Kurgans and Horses)==== |
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Theories favoring the second category of these migrations have the additional attraction to some authors that they would seem to link R1a to well-known language dispersals which resulted in the development of the modern Indo-Aryan language family in India, Central Asia, and the Middle East. In particular, this scenario was linked to the "Kurgan hypothesis" concerning these languages, and these accounts were therefore linked to theories that R1a largely dispersed from Europe (or at least the southeastern edge of Europe) and moved subsequently to Asia. Publications taking this position between 2000 and 2008 include {{Harvcoltxt|Semino et al.|2000}}, {{Harvcoltxt|Passarino et al.|2001}}, {{Harvcoltxt|Passarino et al.|2002}} and {{Harvcoltxt|Wells|2002}}. |
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Such a [[Bronze Age]] European origin for R1a1 in at least parts of Asia has also been argued on the basis of a 2009 study of DNA results from [[Andronovo culture]] remains in South Siberia. The Y DNA was almost exclusively R1a1<ref>{{Harvcoltxt|Keyser et al.|2009}}</ref> This [[archaeological culture]], has also been genetically studied in [[Kazakhstan]], and is thought to have been a carrier of an Indo-Iranian language (the same family of languages as is commonly associated with R1a in modern India) from the direction of Europe. (In particular it has been noted that their mitochondrial DNA is almost entirely of types associated with Europe, and that this Asian population appears to have had a relatively high level of red and blonde hair and blue eyes.)<ref>{{Harvcoltxt|Lalueza-Fox et al.|2004}}</ref> |
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Evidence that during and before the Bronze Age R1a was common in European areas to the west of its modern core range, and even west of the Balkans, has come from ancient samples, which appear to show that R1a was common in this region well before Slavic languages are thought to have arrived.<ref>{{Harvcoltxt|Schilz|2006}}</ref><ref name=Haak2008/><ref>{{Harvcoltxt|Bouakaze et al.|2007}}</ref> |
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====Historic era (Slavic languages): Movements within Europe==== |
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The spread of Slavic peoples and languages might have played a role in further increasing the frequency of R1a1 in parts of Europe, but if so then by all age estimates this would have been on sub-clades of R1a1. So this is not an explanation of the origins and dispersal of R1a1 as a whole. |
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{{Harvcoltxt|Luca et al.|2006}}, looking at SNP and STR markers occurring in the Czech Republic suggested there was evidence for a rapid demographic expansion beginning about 60 to 80 generations ago, which would equate to about 1500 years ago (approx. 500 AD) to 2000 years ago (approx. 1 AD) with a generation time of 25 years. Similar results have been found in Lithuania.{{Citation needed|date=November 2009}} {{Harvcoltxt|Rebala et al.|2007}} also detected Y-STR evidence of a recent Slavic expansion from the area of modern [[Ukraine]]. |
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===South Asian Origin Theories=== |
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Several other studies suggest R1a lineages may have their origins in [[North India]] <ref name=Sharma2007/><ref name=Sengupta2005>{{Harvcoltxt|Sengupta et al.|2005}}</ref><ref name=Sahoo2006>{{Harvcoltxt|Sahoo et al.|2006}}</ref>. |
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As more data has been collated, an increasing number of studies have found South Asia to have the highest diversity of [[microsatellite]] [[Y-STR]] variation, making it likely that South Asia is the original point of dispersal of R1a. Studies which have argued this case most strongly include {{Harvcoltxt|Sengupta et al.|2005}}, {{Harvcoltxt|Sahoo et al.|2006}}, and {{Harvcoltxt|Sharma et al.|2009}}. Studies which have concluded that the data is at least consistent with this scenario include {{Harvcoltxt|Kivisild et al.|2003}}, {{Harvcoltxt|Mirabal et al.|2009}} and {{Harvcoltxt|Underhill et all.|2009}}. The latter two articles, being the most recent and comprehensive, both make the case for Asian origins of R1a the strongest amongst the various possibilities as of late 2009. |
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A particular interest has been taken in investigating the long-presumed connection between [[Indo-Aryan]] origins and higher caste [[Brahmin]]s.<ref>For example {{Harvcoltxt|Wells et al.|2001}}, noted that the Indo-European-speaking Sourashtrans, a population from Tamil Nadu in southern India, have a much higher frequency of M17 [R1a1] than their Dravidian-speaking neighbours, the Yadhavas and Kallars, adding to the evidence that M17 [R1a1] is a diagnostic Indo-Iranian marker.</ref> On the other hand, some authors have not accepted this association.<ref>For example {{Harvcoltxt|Saha et al.|2005}} examined R1a1 in South Indian tribals and Dravidian population groups more closely, and questioned this concept. Their analyses of the haplogroups "indicated no single origin from any lineage but a result of a conglomeration of different lineages from time to time. The phylogenetic analyses indicate a high degree of population admixture and a greater genetic proximity for the studied population groups when compared with other world populations". {{Harvcoltxt|Sharma et al.|2009}} collated information for 2809 Indians (681 Brahmins, and 2128 Tribals and schedule castes). The results showed "no consistent pattern of the exclusive presence and distribution of Y-haplogroups to distinguish the higher-most caste, [[Brahmin]]s, from the lower-most ones, schedule castes and tribals". Brahmins from [[West Bengal]] showed the highest frequency (72.22%) of Y-haplogroups R1a1* hinting that it may have been a founder lineage for this caste group. The authors found it significant that the [[Saharia]] tribe of [[Madhya Pradesh]] had not only 28.07% R1a1, but also 22.8% R1a*, out of 57 people, with such a high percentage of R1a* never having been found before. Based on STR variance the estimated age of R1a* in India was 18,478 years, and for R1a1 it was 13,768 years. In its conclusions, the study proposed "the autochthonous origin and tribal links of Indian Brahmins" as well as the origin of R1a1* in the Indian subcontinent. Chaubey et al. draw the same conclusion that both caste and tribal populations are autochthonous to India. (Chaubey G, Metspalu M, Kivisild T. et al., Peopling of South Asia: investigating the caste-tribe continuum in India, ''Bioessays'' (Jan 2007)). {{Harvcoltxt|Sengupta et al.|2005}} have confirmed R1a's diverse presence even among Indian tribal and lower castes (the so-called [[dalit (outcaste)|untouchables]]) and populations not part of the caste system.</ref> |
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Age estimation techniques play a role in whether authors accept or reject any connection between [[Indo-Aryan languages]], and R1a. In particular, researchers such as Underhill et al. and Mirabal et al., estimate the dispersal of R1a to have started many thousands of years before conventional estimates of the age of this language family, or it's parent, the [[Indo-European]] family. |
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===South-West Asian Origin Theories=== |
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{{Harvcoltxt|Semino et al.|2000}} proposed that a Middle Eastern origin for R1a should be considered, depending upon the strength of arguments for a [[Middle East]]ern origin for [[Indo-European]] languages. |
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Most recently, {{Harvcoltxt|Underhill et al.|2009}} points out, as did {{Harvcoltxt|Regueiro et al.|2006}}, and {{Harvcoltxt|Kivisild et al.|2003}} that the evidence used to argue for South Asian origins of R1a, does not exclude the possibility of a South-West Asian origin: |
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{{cquote|The most distantly related R1a chromosomes, that is, both R1a* and R1a1* (inset, Figure 1), have been detected at low frequency in [[Europe]], [[Turkey]], [[United Arab Emirates]], [[Caucasus]] and [[Iran]] (Supplementary Table S1<ref>The authors also refer here to their references 14, {{Harvcoltxt|Weale et al.|2001}}, and 41, {{Harvcoltxt|Regueiro et al.|2006}}]</ref>). The highest STR diversity of R1a1a*(xM458) chromosomes are observed outside Europe, in particular in South Asia (Figure 1, Supplementary Table S4), but given the lack of informative SNP markers the ultimate source area of haplogroup R1a dispersals remains yet to be refined.}} |
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{{Harvcoltxt|Regueiro et al.|2006}} specifically suggested there was a case for an origin near Southern [[Iran]]. |
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==Popular science== |
==Popular science== |
Revision as of 05:27, 11 November 2009
Haplogroup R1a | |
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Possible time of origin | 30,000-10,000 years BP |
Possible place of origin | proposals include southern Central Asia or Eastern Europe or South Asia |
Ancestor | R1 |
Defining mutations | SRY1532.2 also known as SRY10831.2 has normally been said to define R1a. M17 and M198 define the very dominant sub-clade usually called R1a1. (They always appear together so far.) |
Highest frequencies | Parts of Eastern Europe, Central Asia, and South Asia. (Also found in other parts of Eurasia, except East Asia. See detailed data in article.) |
Haplogroup R1a (R1a) is the name given to a major human Y-chromosome haplogroup within R1 (R-M173). In other words, it is one of the major male-lines of all humanity.
It is found at high frequencies in a wide geographic area extending from South Asia to Central and Eastern Europe and Southern Siberia.[1]
R1a is believed to have originated somewhere within this same area in Eurasia, most likely in the area from Eastern Europe to South Asia. Major reviews of data published in 2009 have lead to a slight preference towards Asia rather than Europe, as will be discussed in detail below.
R1a has a relatively close common ancestry with several other major R haplogroup branches, most notably, R1b, the most common male lineage of Western Europe, and R2, found in South Asia.
Subclades
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2009 latest scheme | |||||||||||||||||||
12009 R1a defining mutations may also include L62 or L63, or these may be redundant with 4 other mutations |
The phylogenetic ("family tree") naming system commonly used for this haplogroup remains inconsistent in different published sources. Although it has not yet used much in published surveys, a fuller survey of known mutations is listed by ISOGG, and also given in Underhill et al. (2009).[2]
Prior to 2009 the mutation SRY1532.2 (or SRY10831.2) defined R1a. However R1a* existed which did not have this mutation but had other defining mutations of basal R1a, M420, M449, M511 and M513 (and/or L62 and L63). Henceforth SRY1532.2 variant clade was moved R1a1 and those mutants that define R1a1 were moved into R1a1a. Underhill et al. (2009) also defines the nascent R1a1 clade with M448, M459, and M516, SNPs. The pre-2009 R1a1 clade which was defined by M17 and M198 with the new name R1a1a had been furhter populated with 5 new SNPs: M417, M512, M514, M515, Page07. This renaming is not universally used and the old and new clades are compared on the right.
With the old nomenclature, the most common clade R1a* with the new nomenclature is now broken down into two rare groups of mutations: R1a*, and R1a1* together representing ~1% of all R1a typed Y chromosomes. The most common clade of R1a is therefore R1a1, and the most common clade of R1a1 is R1a1a. Sharma et al. (2009) found 22.8% R1a*, out of 57 people tested from the Saharia tribe of Madhya Pradesh. Other examples include Northern Iran (1 case out of 33 tests), Crete (1 case of R1a(xM198) out of 193 tests), and Greek Macedonia.[3][4][5]
Distribution
R1a has been found in high frequency at both the eastern and western ends of its core range, for example in some parts of India and Tajikistan on the one hand, and Poland on the other.
Central and Northern Asia
R1a frequencies vary widely between populations within central and northern parts of Eurasia, but R1a is found in areas including Western China and Eastern Siberia. This big variation is possibly a consequence of population bottlenecks in isolated areas and/or the large movements of Turco-Mongols during the historic period. For example, exceptionally high frequencies of R1a1 (R-M17 or R-M198; 50 to 70%) are found among the Ishkashimis, Khojant Tajiks, Kyrgyzs, and in several peoples of Russia's Altai Republic.[6][7][8] Although levels are comparatively low amongst some Turkic-speaking groups (e.g. Turks, Azeris, Kazakhs, Yakuts), levels are very high in certain Turkic- or Mongolic-speaking groups of Northwestern China, such as the Bonan, Dongxiang, Salar, and Uyghurs.[6][9][10] R1a lines propogated north-eastward and are scattered amoung certain indigenous Eastern Siberians, including:Kamchatkans and Chukotkans, and peaking in Itel'man at 22%.[11]
South Asia
In South Asia high levels have been observed in some populations. For example, in the eastern and northern parts of India, among the high caste Bengalis from West Bengal like Brahmins and Kshatriyas (72%), Uttar Pradesh Brahmins (67%), Bihar Brahmins (60%), Punjab (47%), and Gujarat (33%) of male lineages[12] have been observed in this lineage. It is also found in relatively high frequencies in several South Indian Dravidian-speaking tribes including the Chenchu and Valmikis of Andhra Pradesh and the Kallar of Tamil Nadu suggesting that M17 is widespread in tribal southern Indians[13].
South-West Asia
The M17 marker is found in five to ten percent of Middle Eastern men. This is true even in western Iranian populations where Persian, a major Indo-European language with close relatives in high frequency areas in Central and South Asia, is spoken. However, on the Eastern side of Iran, around 35% of men carry the M17 maker.[14] Wells et al. (2001) suggest that the deserts of central Iran acted as "significant barriers to gene flow," and propose two possibilities. First, that the Mesopotamian civilization was densely population relative to immigrating Indo-Iranians; and second, Indo-Iranian languages may have become the predominant language of all Steppe nomads of various ethnic origins. Regueiro et al. (2006), in a study of Iran, noted much higher frequencies in the south than the north and suggested "the lineage may have had an influence on the populations south of the Iranian deserts and where the Dash-e Lut desert would have played a significant role in preventing the expansion of this marker to the north of Iran". The authors suggested that R1a must have originally arrived there prior to any Kurgan/Indo-European expansion into the area, and that the R haplogroup as a whole including R1a may even have roots near Iran.
Europe
In Europe, R1a is found at highest levels among peoples of Eastern European descent (Sorbs, Poles, Russians and Ukranians; 50 to 65%).[15][16][17] Levels in Hungarians have been noted between 20 and 60% [18] The Balkans shows lower frequencies, and significant variation between areas, for example >30% in Slovenia, Croatia and Greek Macedonia, but <10% in Albania, Kosovo and parts of Greece.[17][19][20]. R1a was present in Europe at least 4600 years ago, as demonstrated by Y-DNA extracted from the remains of three individuals near Eulau, Saxony-Anhalt, Germany, discovered in 2005. The discovery demonstrated the appearance of R1a with Corded Ware culture in Central Europe.[21][22]
There is a significant presence in peoples of Scandinavian descent.[23] In Iceland, for instance, R1a accounts for nearly a quarter of the local male Y-DNA. Vikings and Normans may have carried the R1a lineage westward; accounting for a small presence in the British Isles.[24][25][26][27]
In Southern Europe R1a is not normally common but it is widespread and found in significant pockets. Scozzari et al. (2001) found significant levels in the Pas Valley in Northern Spain, and also the areas of Venice, and Calabria in Italy.
Origins
sp |
|
As of November 2009 R1a is defined by the following mutations: L62?, L63?, M420, M449, M511, M513.
The exact point of origin of R1a's dispersal pattern found today remains the subject of discussion. It presumably originated somewhere in the Eurasian landmass, where it is most commonly found today. There is a broad discussion of the possible origin of the largest subclade within R1a*/R1a1* tiers, R1a1a. Establishing the origin of this haplogroup maybe considerably easier than establishing the place of origin of R1a* since R1a* is relatively scarce.
Because R1a* and R1a1* are spread at low frequencies across a broad area, it is difficult to pinpoint sites of diversity, while many studies point toward R1a1a diversity, it should be remembered that R1a1a arose from a single R1a1 Y chromosome and could have done so anywhere that R1a1 had spread at the time R1a1a evolved. Sharma et al. (2009) indicated a site of basal R1a diversity around Bhopal, India with 13 individuals bearing the R1a* mutation, however unlike other study subjects, these R1a were not included as part of the more comprehensive study of Underhill et al. (2009). Mirabal et al. (2009), and Underhill et al. (2009), concluded that there are two separate "poles of the expansion" with similar ages, with South Asian R1a older than European R1a. Of 117 R1a examined from India, none where R1a* or R1a1* (all were R1a1a) contradicting Sharma et al. 2009.
Popular science
Bryan Sykes in his book Blood of the Isles gives (from his imagination) the populations associated with R1a in Europe the name of Sigurd for a clan patriarch, much as he did for mitochondrial haplogroups in his work The Seven Daughters of Eve.
See also
- Human Y-chromosome DNA haplogroups
- Genetics and Archaeogenetics of South Asia
- Pole, Hungarian, two good friends
- Y-DNA haplogroups by ethnic groups
- Nordic R1a Y-DNA Project
Notes
- ^ Underhill et al. (2009)
- ^ ISOGG phylogenetic tree
- ^ Regueiro et al. (2006)
- ^ King et al. & 2007)
- ^ Battaglia et al. (2008)
- ^ a b Wells et al. (2001)
- ^ Kharkov et al. (2007)
- ^ Tambets et al. (2004)
- ^ Wang et al. (2003)
- ^ Zhou et al. (2007)
- ^ Lell et al. (2002)
- ^ Sharma et al. (2007)
- ^ Kivisild et al. (2003)
- ^ "Atlas of the Human Journey - The Genographic Project". The Genographic Project. Retrieved 2009-10-22.
- ^ Balanovsky et al. (2008)
- ^ Behar et al. (2003)
- ^ a b Semino et al. (2000)
- ^ Semino et al. (2000) found a level of 60% but a later study, Tambets et al. (2004), found haplogroup R1a Y-DNA in only 20.4% of a sample of 113 Hungarians. Rosser et al.2000 found haplogroup R1a1-SRY1532b in approximately 22% (8/36) of a Hungarian sample. Battaglia et al. (2008) found haplogroup R1a1a-M17 in approximately 57% of a sample of 53 Hungarians.
- ^ Rosser et al. (2000)
- ^ Pericic et al. (2005)
- ^ Haak et al. (2008)
- ^ The Ysearch number for the Eulau remains is 2C46S.
- ^ Bowden et al. (2008)
- ^ Irish Heritage DNA Project, R1 and R1a
- ^ Passarino et al. (2002)
- ^ Capelli et al. (2003)
- ^ Garvey, D. "Y Haplogroup R1a1". Retrieved 2007-04-23.
References
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