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Haplogroup R1a
Possible time of origin	more recent than 18,000 years BP
Possible place of origin	Asia, probably South Asia. Other possibilities include Central Asia, Middle East, and Eastern Europe.
Ancestor	R1 (R-M173)
Descendants	The most significant sub-clade defined by SNP so far is R-M458 (R1a1a7 in Underhill et al. (2009)).
Defining mutations	1. M420 defines the broadest definition of R1a, and the most recent. 2. Within R-M420, SRY1532.2 also known as SRY10831.2, was said to define R1a until the discovery of M420. 3. M17 and M198 define by far the most dominant sub-clade with R1a generally, referred to as R1a1a since the discovery of M420. (M17 and M198 always appear together so far, and are considered equivalent.)
Highest frequencies	Parts of Eastern Europe, Central Asia, and South Asia. (Also found in other parts of Eurasia, except East Asia. See List of R1a frequency by population)

Haplogroup R1a is the name given to a major human Y-chromosome haplogroup within R1 (also known as R-M173). In other words, it represents one of the major male-lines of humanity.

It is found at high frequencies in a large region extending from South Asia to Central and Southern Siberia.

R1a is believed to have originated somewhere within this same area in Eurasia, most likely in the area from Eastern Europe to South Asia.

Phylogeny (Family Tree)

The most recent publications on this subject have increased knowledge of the complexity of the R1a. The most commonly found type of R1a, now known as R1a1a, represents only one branch of a bigger "family tree". Each major branching of this tree is identified by a corresponding set of known SNP mutations which can be used to test individuals as well as the links between branches as more information becomes available.

Roots of R1a

R1a evolved from a male-line ancestor who was in haplogroup R1, defined by SNP mutation M343. R1a lineages are further distinguished from other surviving R1 lineages by several unique SNP mutations such as M420. (It is therefore the sister clade of R1b - another common R1 lineage, but defined by the M343 mutation.) There is no simple consensus concerning the places in Eurasia where R1, R1a or R1b evolved, although Underhill et al. (2009) recently suggested that "the most distantly related R1a chromosomes, have been detected at low frequency in Europe, Turkey, United Arab Emirates, Caucasus and Iran".

Different meanings of "R1a"

R1 and R1a are "phylogenetic" names, names designed to show a position in a family tree. Names of SNP mutations are also used to name haplogroups. For example, the mutation called "M173" currently identifies R1. Thus R1 can also be called "R-M173". (So when a new branching in a tree is discovered, phylogenetic names must change, while the SNP names remain the same.) There are also "paraclades" which are unknown sections of the tree, whose branching structure is not known. For example R1* men have mutation M173, but they have no other mutations which are yet used to identify branches of R1 (neither M420 nor M343 for example).

The naming system commonly used for this haplogroup remains inconsistent in different published sources. Although it has not yet used much in published surveys, a new tree structure started to be used in 2009. Prior to 2009 the mutation SRY1532.2 was commonly used to identify "R1a". This is also how the term R1a is most often used in publications well into 2009. However the term "R1a" is now increasingly used to refer to a broader family. This includes the "old" R1a but with the addition now of all other lineages which have the M420 mutation. In this newer system, the clade defined by SRY1532.2 moves from "R1a" to "R1a1".

The family tree of R1a as a whole can be divided into three levels of branching. The following summary is based upon the large survey of Underhill et al. (2009) as follows, using the newest phylogenetic naming system:

R1a (R-M420)

This is the broadest definition of R1a, defined by the mutation M420 for example. It is known to have at least two branches: R1a1 (discussed further below), which makes up the vast majority, and R1a*, the paraclade, made up of an unknown numbers of branches. R1a* in this case is defined as M420 positive but SRY1532.2 negative. (Such cases would have been classified as R-M173* before the discovery of M420.)

Underhill et al. (2009) found only isolated samples of the paraclade, apparently mostly in the Middle East and Caucasus: 1/121 Omanis, 2/150 Iranians, 1/164 in the United Arab Emirates, 3/612 in Turkey. 7224 more tests in 73 other Eurasian population showed no sign of this category so far. Mutations understood to be equivalent to M420 include M449, M511, M513, L62, and L63.

R-SRY1532.2

R1a1 is currently defined by SRY1532.2, also referred to as SRY10831.2. This family of lineages is dominated by very large and well-defined branch R-M17), positive for M17 and M198 (see below). The paraclade R1a1* (old R1a*) is in this case SRY1532.2 positive, but negative for M17 and/or M198.

Underhill et al. (2009) again found only limited examples of the paraclade, looking at many different surveys. However it does appear to be spread over a wider geographical range in Eurasia than the R1a* paraclade discussed above: 1/51 in Norway, 3/305 in Sweden, 1/57 Greek Macedonians, 1/150 Iranians, 2/734 Ethnic Armenians, 1/141 Kabardians. Sharma et al. (2009) also found 13/57 people tested from the Saharia tribe of Madhya Pradesh, and 2/51 amongst Kashmir Pandits. SNP mutations understood to be always occurring with SRY1532.2 include M448, M459, and M516.

R-M17/R-M198

M17 or M198 currently define as R1a1a (old R1a1) each used as a marker identifying the same clade in different publications. These two SNPs always appear together. R1a1a is the dominant within the R-M420 clade, and most statistical or other analysis is by definition focused upon it. This clade also has some sub-clades of its own, although a large proportion of R-M17/R-M198 has however not yet been categorized into branches defined by mutations, and is therefore referred to as R1a1* (old nomenclature) or R1a1a* (new nomenclature). SNP mutations understood to be always occurring with M17 and M198 include M417, M512, M514, M515, and rs34297606.

R1a1a subclades

So far, 8 sub-clades of R1a1a are known, R1a1a1 to R1a1a8. However only three subclades are so far known to have significant frequencies, R1a1a3, R1a1a6 and R1a1a7. R1a1a3 was first identified as R1a1c (old nomenclature) and is defined by the M64.2, M87, and M204 SNPs is apparently rare, found in 1 of 117 males typed in southern Iran. R1a1a6, defined by M434, was announced in Underhill et al. (2009). M434 was detected in 14 people (of how many tested) from Pakistan to Oman and is likely to reflect a recent mutation that took place in the area of Pakistan.

The largest defined subclade of R1a1a is R1a1a7. R1a1a7 is defined by M458 and was first described in Underhill et al. (2009). M458 is found primarily Europe (What about north America?), with its highest frequencies in Central and Southern Poland. R1a1a7 has its own subclade, defined by the M334 marker.

Distribution of R1a1a (R-M17 or R-M198), the dominant sub-clade of R1a

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. Throughout all of these regions, R1a is dominated by the R1a1a (R-M17 or R-M198) sub-clade.

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. 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. R1a lines propogated north-eastward and are scattered amoung certain indigenous Eastern Siberians, including:Kamchatkans and Chukotkans, and peaking in Itel'man at 22%.

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 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.

Western Asia

The M17 marker is present all around Western Asia, in widely varying concentrations, from almost no presence in areas such as Jordan, to much higher levels in parts of Turkey and Iran.

Nasidze et al. (2004) found R1a in approximately 20% of Iranian males from the cities of Tehran and Isfahan. 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 of south of Iran and the Dash-e Lut (sic.) desert would have played a significant role in preventing the expansion of this marker to the north of Iran".

Europe

In Europe, R1a, again almost entirely in the M17/M198 sub-clade, is found at highest levels among peoples of Eastern European descent (Sorbs, Poles, Russians and Ukranians; 50 to 65%). Levels in Hungarians have been noted between 20 and 60% 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.. In the Baltic countries R1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).

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.

There is a significant presence in peoples of Scandinavian descent. 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.

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 and hypothesized migrations of R1a1a (R-M17/R-M198)

Most recent discussions of R1a origins concern the dominant R1a1a (M17/M198) sub-clade. There are two foci of high frequency of R1a1a (the majority of R1a), one in South Asia, near North India, and the other in Eastern Europe, in the area of the Ukraine. On the one hand, the highest frequency level observed in any large population so far has been found in some South Asian groups. On the other hand, until 2009 claims regarding the oldest R1a bearing populations varied greatly between different articles. Some of the older articles supported origins in Central Asia and Europe. In 2009, two large studies of available data, 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 R1a1a (M17/M198) older than European R1a1a. Both articles felt the data to be most consistent with Asian, as opposed to European origins for R1a1a, and in particularly, South Asian origins.(Mirabal et al. (2009) additionally felt the data to be consistent with central Asian, while Underhill et al. (2009) took to the data to be consistent with Western Asian origins.)

Central Asian origin proposal

Cordaux et al. (2004) argued, Citing data from 3 earlier publications, that R-M17 (R1a1a) Y chromosomes most probably have a central Asian origin. Central Asia is still considered a possible place of origin by Mirabal et al. (2009) after their larger analysis of more recent data. However these authors do not clearly distinguish the case being made for Central Asia for the case being made for Asia, particularly South Asia, more generally.

Eastern European migration hypotheses

Theories that the earliest generations of R1a1a (M17/M198) men originated in Eastern Europe have become less common with the publication of bigger and more international surveys. However suggestions have been made which associate the distribution of R1a clades with several proposed movements of people in history and prehistory in Eastern Europe. As usual, these suggestions mainly concern the R1a1a sub-clade defined by M17/M198, because this is the dominant R1a clade, and the only one for which there is significant data.

• The period from the end of the Ice Age until the Mesolithic. The spread from a Ukrainian refugium during the Late Glacial Maximum.
• The European Neolithic, for example the LBK (Linear Pottery) Culture and Corded Ware Culture.
• 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).
• Historical Era. The spread of Slavic languages and migrations in late Classical times.

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.

Europe from the end of the Ice Age until the Mesolithic

Underhill et al. (2009) estimated that R1a1a (M17/M198) dispersed in parts of Europe approximately 11,000 years ago, prior to the Neolithic period. Age estimates of this depth for R1a1a come from papers using the methodology described by Zhivotovsky et al. (2004), the latest such example being Mirabal et al. (2009) and Underhill et al. (2009). (Other methods tend to give much younger estimates for any given set of data.) Researchers using this estimation method therefore believe any Bronze Age or more recent dispersals affecting modern R1a1a diversity can not involve the clade as a whole, but only some branches.

Neolithic

Underhill et al. (2009) associates the Neolithic period in Europe with the R-M458 sub-clade, as announced in that article, specifically with the Corded Ware Horizon. Other authors have for example linked R1a haplotypes in Norway with "the spread of the Corded Ware and Battle-Axe cultures from central and east Europe".

Once again, it should be noted that this is using the Zhivitovsky age estimation method. Other methods would suggest that all or most R1a in Europe may have only arrived at a time close to the Neolithic.

Bronze Age (Indo Europeans, Indo-Aryans, Kurgans and horses)

Proposals of Bronze Age R1a migrations have the attraction to some authors that they would seem to link R1a1a (M17/M198) 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. This popular scenario has been linked to the "Kurgan hypothesis" concerning the origin of these languages. Making this link therefore involves assuming that Asian R1a, or at least a large segment of it, dispersed from Europe, or at least from the Eurasian Steppe which protrudes into the southeastern edge of Europe.

Such a Bronze Age European origin for R1a1a 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 R1a of some type. This archaeological culture, has also been genetically studied in Kazakhstan, and is thought to have been a carrier of an Indo-Aryan 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.)

Evidence that during and before the Bronze Age R1a existed in Europe 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. This was probably R1a1a* (M17/M198 positive, M458 negative) according to Underhill et al. (2009).

European migrations within the Historic Era

The spread of Slavic peoples and languages might have played a role in further increasing the frequency of R1a1a (M17/M198) in parts of Europe, but if so then by all age estimates this would have been after R1a1 had already dispersed as widely as both Central Europe and India. So this is not an explanation of the origins and dispersal of R1a1 as a whole.

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. Rebala et al. (2007) also detected Y-STR evidence of a recent Slavic expansion from the area of modern Ukraine. This evidence corresponds to population movements during the late Classical Migration Period.

South Asian origin hypothesis

Several other studies suggest R1a lineages generally may have their origins in South Asia .

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 within R1a1a (M17/M198), making it likely that South Asia is the original point of dispersal. Studies which have argued this case most strongly include Sengupta et al. (2005), Sahoo et al. (2006), and Sharma et al. (2009). Studies which have concluded that the data is at least consistent with this scenario include Kivisild et al. (2003), Mirabal et al. (2009) and Underhill et all. (2009) harvcoltxt error: no target: CITEREFUnderhill_et_all.2009 (help). The latter two articles, being the most recent and comprehensive, both make the case for Asian origins of R1a1a the strongest amongst the various possibilities as of late 2009.

A particular interest has been taken in investigating the long-presumed connection between Indo-Aryan origins and higher caste Brahmins. On the other hand, some authors have not accepted this association.

Age estimation techniques play a role in whether authors accept or reject any connection between Indo-Aryan languages, and R1a in any broad sense. In particular, researchers such as Underhill et al. and Mirabal et al., estimate the dispersal of R1a1a (M17/M198) to be much older than the Indo-Aryan language family.

Middle Eastern origin hypothesis

As mentioned above, R1a haplotypes are less common in most of the Middle East than they are in either South Asia or Eastern Europe or much of Central Asia. It has nevertheless been mentioned in speculation about the origins of the clade, bother because there are interesting pockets of high frequency, for example in some parts of Iran and amongst some Kurdish populations, and also because the rarer branches of R1a (R1a*, R1a1*) are less rare in some of these regions.

Semino et al. (2000) proposed that a Middle Eastern origin for R1a should be considered, depending upon the strength of arguments for a Middle Eastern origin for Indo-European languages. However, Nasidze et al. (2004) 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.

Most recently, Underhill et al. (2009) points out, as did Regueiro et al. (2006), and 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:

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). 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.

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

• List of R1a frequency by population
• Human Y-chromosome DNA haplogroups
• Genetics and Archaeogenetics of South Asia
• Y-DNA haplogroups by ethnic groups
• Nordic R1a Y-DNA Project
• Somerled

v t e Phylogenetic tree of human Y-chromosome DNA haplogroups
This article needs to be updated. Please help update this article to reflect recent events or newly available information. (February 2021)
"Y-chromosomal Adam" A00 A0-T  A0 A1  A1a A1b A1b1 BT B CT DE CF D E C F F1  F-Y27277   F3  GHIJK G HIJK IJK H IJ K I   J     LT        K2  I1   I2  J1   J2  L     T  K2e K2d K2c K2b   K2a K2b1    P  K-M2313  S   M     P1   NO1 P1c P1b P1a N O R Q
Y-DNA by population Y-DNA haplogroups of historic people Footnotes Van Oven M, Van Geystelen A, Kayser M, Decorte R, Larmuseau HD (2014). "Seeing the wood for the trees: a minimal reference phylogeny for the human Y chromosome". Human Mutation. 35 (2): 187–91. doi:10.1002/humu.22468. PMID 24166809. S2CID 23291764. International Society of Genetic Genealogy (ISOGG; 2015), Y-DNA Haplogroup Tree 2015. (Access date: 1 February 2015.) Haplogroup A0-T is also known as A-L1085 (and previously as A0'1'2'3'4). Haplogroup A1 is also known as A1'2'3'4. F-Y27277, sometimes known as F2'4, is both the parent clade of F2 and F4 and a child of F-M89. Haplogroup LT (L298/P326) is also known as Haplogroup K1. Between 2002 and 2008, Haplogroup T-M184 was known as "Haplogroup K2". That name has since been re-assigned to K-M526, the sibling of Haplogroup LT. Haplogroup K2b (M1221/P331/PF5911) is also known as Haplogroup MPS. Haplogroup K2b1 (P397/P399) is also known as Haplogroup MS, but has a broader and more complex internal structure. Haplogroup P (P295) is also klnown as K2b2. K-M2313*, which as yet has no phylogenetic name, has been documented in two living individuals, who have ethnic ties to India and South East Asia. In addition, K-Y28299, which appears to be a primary branch of K-M2313, has been found in three living individuals from India. See: Poznik op. cit.; YFull YTree v5.08, 2017, "K-M2335", and; PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (Access date of these pages: 9 December 2017) Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.) Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)

Notes

1. Karafet et al. (2008)
2. ^ Underhill et al. (2009)
3. Underhill et al. (2009)
4. Underhill et al. (2009)
5. ISOGG phylogenetic tree
6. SRY1532.2 is also known as SRY10831.2
7. ISOGG phylogeny webpage 2009
8. Regueiro et al. (2006)
9. ^ Wells et al. (2001)
10. Kharkov et al. (2007)
11. Tambets et al. (2004)
12. Wang et al. (2003)
13. Zhou et al. (2007)
14. Lell et al. (2002)
15. ^ Sharma et al. (2007)
16. Kivisild et al. (2003)
17. Flores et al. (2005) harvcoltxt error: no target: CITEREFFlores_et_al.2005 (help)
18. Nasidze et al. (2004)
19. Nasidze et al. (2005)
20. Balanovsky et al. (2008)
21. Behar et al. (2003)
22. ^ Semino et al. (2000)
23. 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 SRY1532b positive lineages 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.
24. Rosser et al. (2000)
25. Pericic et al. (2005)
26. Kasperaviciūte et al. (2005)
27. ^ Haak et al. (2008)
28. The Ysearch number for the Eulau remains is 2C46S.
29. Bowden et al. (2008)
30. Irish Heritage DNA Project, R1 and R1a
31. Passarino et al. (2002)
32. Capelli et al. (2003)
33. Garvey, D. "Y Haplogroup R1a1". Retrieved 2007-04-23.
34. Sharma et al. (2009)
35. Wells et al. (2001), Semino et al. (2000), and Quintana-Murci et al. (2001)
36. Semino et al. (2000) proposed quite early that there may have been two expansions, suggesting that the spread of R1a from 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, 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.
37. Dupuy et al. (2005)
38. see: Semino et al. (2000), Passarino et al. (2001), Passarino et al. (2002) and Wells (2002)
39. Keyser et al. (2009)
40. Lalueza-Fox et al. (2004)
41. Schilz (2006) harvcoltxt error: no target: CITEREFSchilz2006 (help)
42. Bouakaze et al. (2007)
43. Sengupta et al. (2005)
44. Sahoo et al. (2006)
45. For example 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 than their Dravidian-speaking neighbours, the Yadhavas and Kallars, adding to the evidence that M17 is a diagnostic Indo-Iranian marker.
46. For example 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". 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, Brahmins, 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)). Sengupta et al. (2005) have confirmed R1a's diverse presence even among Indian tribal and lower castes (the so-called untouchables) and populations not part of the caste system.
47. The authors also refer here to their references 14, Weale et al. (2001), and 41, Regueiro et al. (2006)]

References

• Adams; et al. (2008), "The Genetic Legacy of Religious Diversity and Intolerance: Paternal Lineages of Christians, Jews, and Muslims in the Iberian Peninsula", The American Journal of Human Genetics, 83: 725, doi:10.1016/j.ajhg.2008.11.007 {{citation}}: Explicit use of et al. in: |author= (help)
• Al Zahery; et al. (doi:10.1016/S1055-7903(03)00039-3), "Y-chromosome and mtDNA polymorphisms in Iraq, a crossroad of the early human dispersal and of post-Neolithic migrations" (PDF), Molecular Phylogenetics and Evolution, doi:10.1016/S1055-7903(03)00039-3 {{citation}}: Check date values in: |year= (help); Explicit use of et al. in: |author= (help)CS1 maint: year (link)
• Balanovsky; et al. (2008), "Two Sources of the Russian Patrilineal Heritage in Their Eurasian Context", AJHG, 82 (1): 236–250, doi:10.1016/j.ajhg.2007.09.019 {{citation}}: Explicit use of et al. in: |last= (help)
• Bamshad; et al. (2001), "Genetic evidence on the origins of Indian caste populations", Genome Research, vol. 11, no. 6, pp. 994–1004, doi:10.1101/gr.GR-1733RR, PMID 11381027 {{citation}}: Explicit use of et al. in: |author= (help).
• Barać L, Pericić M, Klarić IM; et al. (2003). "Y chromosomal heritage of Croatian population and its island isolates" (PDF). Eur. J. Hum. Genet. 11 (7): 535–42. doi:10.1038/sj.ejhg.5200992. PMID 12825075. {{cite journal}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
• Battaglia; et al. (2008), "Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe", European Journal of Human Genetics, doi:10.1038/ejhg.2008.249 {{citation}}: Explicit use of et al. in: |author= (help)
• Behar; et al. (2003), "Multiple Origins of Ashkenazi Levites: Y Chromosome Evidence for Both Near Eastern and European Ancestries" ( – ), Am. J. Hum. Genet., vol. 73, pp. 768–779, doi:10.1086/378506, PMID 13680527 {{citation}}: Explicit use of et al. in: |author= (help); External link in |format= (help). Also at http://www.ucl.ac.uk/tcga/tcgapdf/Behar-AJHG-03.pdf and http://www.familytreedna.com/pdf/400971.pdf
• Bouakaze; et al. (2007), "First successful assay of Y-SNP typing by SNaPshot minisequencing on ancient DNA", International Journal of Legal Medicine, 121 (6): 493–9 {{citation}}: Explicit use of et al. in: |author= (help)
• Bowden; et al. (2008), "Excavating Past Population Structures by Surname-Based Sampling: The Genetic Legacy of the Vikings in Northwest England", Molecular Biology and Evolution, 25 (2): 301–309, doi:10.1093/molbev/msm255 {{citation}}: Explicit use of et al. in: |author= (help)
• Capelli; et al. (2003), "A Y Chromosome Census of the British Isles", Current Biology, 13 (11): 979–84, doi:10.1016/S0960-9822(03)00373-7 {{citation}}: Explicit use of et al. in: |last= (help) also at
• Cinnioğlu; et al. (2004), "Excavating Y-chromosome haplotype strata in Anatolia" (PDF), Hum Genet, 114: 127, doi:10.1007/s00439-003-1031-4 {{citation}}: Explicit use of et al. in: |last= (help)
• Cordaux; et al. (2004), "Independent Origins of Indian Caste and Tribal Paternal Lineages", Current Biology: 231–235, doi:10.1016/j.cub.2004.01.024 {{citation}}: Explicit use of et al. in: |author= (help); Unknown parameter |vol= ignored (|volume= suggested) (help)
• Dupuy; et al. (2005), "Geographical heterogeneity of Y-chromosomal lineages in Norway" (PDF), Forensic Science International, 164: 10–19 {{citation}}: Explicit use of et al. in: |author= (help)
• Firasat; et al. (2006), "Y-chromosomal evidence for a limited Greek contribution to the Pathan population of Pakistan", European Journal of Human Genetics, 15: 121–126, doi:10.1038/sj.ejhg.5201726 {{citation}}: Explicit use of et al. in: |author= (help)
• Fornarino; et al. (2009), "Mitochondrial and Y-chromosome diversity of the Tharus (Nepal): a reservoir of genetic variation", BMC Evolutionary Biology (154), doi:10.1186/1471-2148-9-154 {{citation}}: Explicit use of et al. in: |author= (help); Unknown parameter |vol= ignored (|volume= suggested) (help)CS1 maint: unflagged free DOI (link)
• Gimbutas (1970), Indo-European and Indo-Europeans, Univ. of Pennsylvania Press, Philadelphia, PA, pp. 155–195
• Haak; et al. (2008), Ancient DNA, Strontium isotopes, and osteological analyses shed light on social and kinship organization of the Later Stone Age, doi:10.1073/pnas.0807592105 {{citation}}: Explicit use of et al. in: |author= (help); Unknown parameter |vol= ignored (|volume= suggested) (help)
• Helgason; et al. (2000), "Estimating Scandinavian and Gaelic Ancestry in the Male Settlers of Iceland", Am Journal of Human Genetics, doi:10.1086/303046 {{citation}}: Explicit use of et al. in: |author= (help)
• Karafet; et al. (2008), Abstract "New Binary Polymorphisms Reshape and Increase Resolution of the Human Y-Chromosomal Haplogroup Tree", Genome Research, 18: 830, doi:10.1101/gr.7172008, PMC 2336805, PMID 18385274 {{citation}}: Check |url= value (help); Explicit use of et al. in: |last= (help); Unknown parameter |month= ignored (help). Published online April 2, 2008. See also Supplementary Material.
• Kasperaviciūte; et al. (2005), "Y Chromosome and Mitochondrial DNA Variation in Lithuanians", Annals of Human Genetics, 68 (5): 438–452, doi:10.1046/j.1529-8817.2003.00119.x {{citation}}: Explicit use of et al. in: |author= (help)
• Kayser; et al. (2005), "Significant genetic differentiation between Poland and Germany follows present-day political borders, as revealed by Y-chromosome analysis", Human Genetics, 117 (5): 428–443, doi:10.1007/s00439-005-1333-9 {{citation}}: Explicit use of et al. in: |last= (help) A copy can be found here.
• Keyser; et al. (2009), Ancient DNA provides new insights into the history of south Siberian Kurgan people {{citation}}: Explicit use of et al. in: |author= (help)
• Kharkov; et al. (2004), "Gene Pool Structure of Eastern Ukrainians as Inferred from the Y-Chromosome Haplogroups", Russian Journal of Genetics, 40(3): 326, doi:10.1023/B:RUGE.0000021635.80528.2f {{citation}}: Explicit use of et al. in: |last= (help) A copy can be found here.
• Kharkov; et al. (2005), "Frequencies of Y Chromosome Binary Haplogroups in Belarussians", Russian Journal of Genetics, 41(8): 928–931, doi:10.1007/s11177-005-0182-x {{citation}}: Explicit use of et al. in: |last= (help) A copy can be found here.
• Kharkov; et al. (2007), "Gene Pool Differences between Northern and Southern Altaians Inferred from the Data on Y-Chromosomal Haplogroups" (PDF), Russian Journal of Genetics, 43 (5), doi:10.1134/S1022795407050110 {{citation}}: Explicit use of et al. in: |author= (help)
• King; et al. (2008), "Differential Y-chromosome Anatolian Influences on the Greek and Cretan Neolithic" (PDF), Annals of Human Genetics, 72: 205–214, doi:10.1111/j.1469-1809.2007.00414.x {{citation}}: Explicit use of et al. in: |last= (help)
• Kivisild; et al. (2003), "The Genetic Heritage of the Earliest Settlers Persists Both in Indian Tribal and Caste Populations", AJHG, vol. 72, p. 313, doi:10.1086/346068 {{citation}}: Explicit use of et al. in: |author= (help).
• Lalueza-Fox; et al. (2004), "Unravelling migrations in the steppe: mitochondrial DNA sequences from ancient central Asians.", Proc Biol Sci., 271 (1542): 941–947 {{citation}}: Explicit use of et al. in: |author= (help); Unknown parameter |PMCID= ignored (|pmc= suggested) (help)
• Lell; et al. (2002), "The Dual Origin and Siberian Affinities of Native American Y Chromosomes" (PDF), Am. J. Hum. Genet., 70: 192–206, doi:10.1086/338457 {{citation}}: Explicit use of et al. in: |last= (help)
• Luca; et al. (2006), "Y-Chromosomal Variation in the Czech Republic", American Journal of Physical Anthropology, 132: 132, doi:10.1002/ajpa.20500 {{citation}}: Explicit use of et al. in: |last= (help)
• Malaspina; et al. (2003), Analysis of Y-chromosome variation in modern populations at the European-Asian border (PDF), p. 309–313 {{citation}}: Explicit use of et al. in: |author= (help) in K. Boyle, C. Renfrew, and M. Levine, eds. Ancient interactions: east and west in Eurasia. McDonald Institute for Archaeological Research Monograph Series, Cambridge University Press, Cambridge
• Marjanovic D, Fornarino S, Montagna S; et al. (2005). "The peopling of modern Bosnia-Herzegovina: Y-chromosome haplogroups in the three main ethnic groups". Ann. Hum. Genet. 69 (Pt 6): 757–63. doi:10.1111/j.1529-8817.2005.00190.x. PMID 16266413. {{cite journal}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
• Mirabal; et al. (2009), "Y-Chromosome distribution within the geo-linguistic landscape of northwestern Russia", European Journal of Human Genetics, 17: 1260–1273, doi:10.1038/ejhg.2009.6 {{citation}}: Explicit use of et al. in: |author= (help)
• Mukherjee; et al. (2001), "High-resolution analysis of Y-chromosomal polymorphisms reveals signatures of population movements from central Asia and West Asia into India", Journal of Genetics, vol. 80, no. 3 (published December, 2001), pp. 125–135, doi:10.1007/BF02717908 {{citation}}: Check date values in: |publication-date= (help); Explicit use of et al. in: |author= (help).
• Nasidze; et al. (2004), "Mitochondrial DNA and Y-Chromosome Variation in the Caucasus" (PDF), Annals of Human Genetics, 68: 205–221, doi:10.1046/j.1529-8817.2004.00092.x {{citation}}: Explicit use of et al. in: |last= (help)
• Nasidze; et al. (2005), "MtDNA and Y-chromosome Variation in Kurdish Groups" (PDF), Annals of Human Genetics, 69: 401–412, doi:10.1046/j.1529-8817.2005.00174.x {{citation}}: Explicit use of et al. in: |author= (help)
• Passarino; et al. (2001), "The 49a,f haplotype 11 is a new marker of the EU19 lineage that traces migrations from northern regions of the black sea", Hum. Immunol., vol. 62, no. 9, pp. 922–932, doi:10.1016/S0198-8859(01)00291-9 {{citation}}: Explicit use of et al. in: |author= (help).
• Passarino; et al. (2002), "Different genetic components in the Norwegian population revealed by the analysis of mtDNA and Y chromosome polymorphisms", Eur. J. Hum. Genet., vol. 10, no. 9, pp. 521–9, doi:10.1038/sj.ejhg.5200834, PMID 12173029 {{citation}}: Explicit use of et al. in: |author= (help).
• Pericic; et al. (2005), "High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations", Mol. Biol. Evol., vol. 22, no. 10, pp. 1964–75, doi:10.1093/molbev/msi185, PMID 15944443 {{citation}}: Explicit use of et al. in: |author= (help).
• Qamar; et al. (2002), "Y-Chromosomal DNA Variation in Pakistan", The American Journal of Human Genetics, doi:10.1086/339929 {{citation}}: Explicit use of et al. in: |author= (help)
• Quintana-Murci; et al. (2001), "Y-chromosome lineages trace diffusion of people and languages in southwestern Asia", Am. J. Hum. Genet: 537–542 {{citation}}: Explicit use of et al. in: |author= (help); Unknown parameter |vol= ignored (|volume= suggested) (help)
• Rebala; et al. (2007), "Y-STR variation among Slavs: evidence for the Slavic homeland in the middle Dnieper basin", Journal of Human Genetics, 52 (5): 406–414, doi:10.1007/s10038-007-0125-6 {{citation}}: Explicit use of et al. in: |author= (help)
• Regueiro; et al. (2006), "Iran: Tricontinental Nexus for Y-Chromosome Driven Migration" (PDF), Hum Hered, 61: 132–143, doi:10.1159/000093774 {{citation}}: Explicit use of et al. in: |last= (help)
• Rosser; et al. (2000), "Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language", American Journal of Human Genetics, 67: 1526–1543., doi:10.1086/316890 {{citation}}: Explicit use of et al. in: |last= (help)
• Saha; et al. (2005), "Genetic affinity among five different population groups in India reflecting a Y-chromosome gene flow", J. Hum. Genet., vol. 50, no. 1, pp. 49–51, doi:10.1007/s10038-004-0219-3, PMID 15611834 {{citation}}: Explicit use of et al. in: |author= (help).
• Sahoo; et al. (2006), "A prehistory of Indian Y chromosomes: Evaluating demic diffusion scenarios", Proceedings of the National Academy of Sciences, 103 (4): 843–848, doi:10.1073/pnas.0507714103 {{citation}}: Explicit use of et al. in: |last= (help)
• Sanchez; et al. (2003), "Multiplex PCR and minisequencing of SNPs—a model with 35 Y chromosome SNPs", Forensic Sci Int, 137: 74–84, doi:10.1016/S0379-0738(03)00299-8 {{citation}}: Explicit use of et al. in: |author= (help)
• Scozzari; et al. (2001), "Human Y-Chromosome Variation in the Western Mediterranean Area: Implications for the Peopling of the Region" (PDF), Human Immunology, 62: 871–884, doi:10.1016/S0198-8859(01)00286-5 {{citation}}: Cite has empty unknown parameter: |unused_data= (help); Explicit use of et al. in: |author= (help)
• Semino; et al. (2000), "The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective" (PDF), Science, vol. 290, pp. 1155–59, doi:10.1126/science.290.5494.1155, PMID 11073453 {{citation}}: Explicit use of et al. in: |author= (help). Copy can be found here.
• Sengupta; et al. (2005), "Polarity and Temporality of High-Resolution Y-Chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influence of Central Asian Pastoralists", Am. J. Hum. Genet., vol. 78, no. 2, pp. 202–21, doi:10.1086/499411, PMID 16400607 {{citation}}: Explicit use of et al. in: |author= (help).
• Sharma; et al. (2007), "The Autochthonous Origin and a Tribal Link of Indian Brahmins: Evaluation Through Molecular Genetic Markers" (PDF), THE AMERICAN SOCIETY OF HUMAN GENETICS 57th Annual Meeting {{citation}}: Explicit use of et al. in: |last= (help)
• Sharma; et al. (2009), "The Indian origin of paternal haplogroup R1a1(*)substantiates the autochthonous origin of Brahmins and the caste system", J. Hum.Genet., 54 (1): 47–55, doi:10.1038/jhg.2008.2, PMID: 19158816 {{citation}}: Explicit use of et al. in: |last= (help)CS1 maint: date and year (link)
• Shilz (2006), Molekulargenetische Verwandtschaftsanalysen am prähistorischen Skelettkollektiv der Lichtensteinhöhle, Dissertation, Göttingen (PDF)
• Tambets; et al. (2004), "The Western and Eastern Roots of the Saami—the Story of Genetic 'Outliers' Told by Mitochondrial DNA and Y Chromosomes", American Journal of Human Genetics, 74: 661–682, doi:10.1086/383203 {{citation}}: Explicit use of et al. in: |last= (help)
• Underhill; et al. (2009), "Separating the post-Glacial coancestry of European and Asian Y chromosomes within haplogroup R1a", European Journal of Human Genetics, doi:doi:10.1038/ejhg.2009.194 {{citation}}: Check |doi= value (help); Explicit use of et al. in: |author= (help)
• Varzari, Alexander (2006), "Population History of the Dniester-Carpathians: Evidence from Alu Insertion and Y-Chromosome Polymorphisms" (PDF), Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität München
• Völgyi; et al. (2008), "Hungarian population data for 11 Y-STR and 49 Y-SNP markers", Forensic Science International: Genetics, 3, doi:10.1016/j.fsigen.2008.04.006 {{citation}}: Explicit use of et al. in: |author= (help)
• Wang; et al. (2003), "The origins and genetic structure of three co-resident Chinese Muslim populations: the Salar, Bo'an and Dongxiang", Human Genetics {{citation}}: Explicit use of et al. in: |last= (help)
• Weale; et al. (2001), "Armenian Y chromosome haplotypes reveal strong regional structure within a single ethno-national group" (PDF), Hum Genet, 109: 659–674, doi:10.1007/s00439-001-0627-9 {{citation}}: Explicit use of et al. in: |author= (help)
• Weale; et al. (2002), "Y Chromosome Evidence for Anglo-Saxon Mass Migration" (PDF), Mol. Biol. Evol., vol. 19, no. 7, pp. 1008–1021, PMID 16400607 {{citation}}: Explicit use of et al. in: |author= (help).
• Wells; et al. (2001), "The Eurasian Heartland: A continental perspective on Y-chromosome diversity", Proc. Natl. Acad. Sci. U. S. A., vol. 98, no. 18, pp. 10244–9, doi:10.1073/pnas.171305098, PMID 11526236 {{citation}}: Explicit use of et al. in: |author= (help). Also at http://www.pnas.org/cgi/reprint/98/18/10244.pdf
• Wells, Spencer (2002), The Journey of Man: A Genetic Odyssey, Princeton University Press.
• Wilson; et al. (2001), "Genetic evidence for different male and female roles during cultural transitions in the British Isles", Proc. Natl. Acad. Sci. USA (9): 5078–5083, doi:10.1073/pnas.071036898 {{citation}}: Explicit use of et al. in: |author= (help); Unknown parameter |volum= ignored (help)
• Zerjal; et al. (2001), "Geographical, linguistic, and cultural influences on genetic diversity: Y-chromosomal distribution in Northern European populations", Mol Biol Evol, 18: 1077–1087 {{citation}}: Explicit use of et al. in: |author= (help)
• Zerjal; et al. (2002), "A Genetic Landscape Reshaped by Recent Events: Y-Chromosomal Insights into Central Asia", Am J Hum Genet., 71 (3): 466–482, doi:10.1086/342096 {{citation}}: Explicit use of et al. in: |last= (help)
• Zhou; et al. (2007), "Testing the hypothesis of an ancient Roman soldier origin of the Liqian people in northwest China: a Y-chromosome perspective", Journal of Human Genetics, 52 (7): 584, doi:10.1007/s10038-007-0155-0 {{citation}}: Explicit use of et al. in: |last= (help)CS1 maint: extra punctuation (link)
• Zhao; et al. (2009), "Presence of three different paternal lineages among North Indians: A study of 560 Y chromosomes", Annals of Human Biology, 36: 1–14, doi:10.1080/03014460802558522 {{citation}}: Explicit use of et al. in: |author= (help)
• Zhivotovsky; et al. (2004), "The effective mutation rate at Y chromosome short tandem repeats, with application to human population-divergence time", Am J Hum Genet, 74: 50–61 {{citation}}: Explicit use of et al. in: |author= (help)

Projects

• FTDNA R1a Y-chromosome Haplogroup Project
• R1a International Y-DNA Project
• Danish Demes Regional DNA Project: Y-DNA Haplogroup R1a
• British Isles DNA Project

• Human Y-DNA haplogroups
• Human genetic history