| Haplogroup R1a | |
| Time of origin | more recent than 18,000 years BP[1] |
| Place of origin | Asia, probably South Asia. Other possibilities include Central Asia, Middle East, and Eastern Europe. |
| Ancestor | R1 (R-M173) |
| Descendants | R1a1a1 to R1a1a8. R-M458 being the most significant (R1a1a7 in Underhill et al. (2009)). |
|---|---|
| Defining mutations | 1. M420 now defines R1a in the broadest sense.[2] 2. Within R1a, SRY1532.2 also known as SRY10831.2, now defines R1a1, previously R1a. 3. M17 and M198 (equivalent to one another) define R1a1a, which is by far the most common type of R1a. |
| 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, a major grouping within all Y-DNA types. As a haplogroup, R1a represents a grouping of Y-chromosome types because of DNA similarities, such as unique mutations. These Y-chromosomes have differences, or markers, that other Y-chromosomes lack. Males that carry the same markers are considered to be related by common descent from a single male ancestor.
R1a is a particularly common category of male line, found especially in a large region extending from South Asia and Southern Siberia to Central Europe and Scandinavia.[2] It is therefore commonly referred to in the fields of population genetics and genetic genealogy.
R1a can be viewed as a family tree of male lines, with branches. However one sub-branch, R1a1a, is much more common than the others. The mutation that is currently used to define the R1a family most broadly is M420. The recent discovery of M420 resulted in a reorganization of what is known about the R1a family tree. In particular, this discovery demonstrated that there are rare relatives of R1a1 which are only found in Asia, including the area of the Middle East, where R1a1a is relatively uncommon.
The origin of R1a is uncertain. However it is believed to have originated somewhere within Eurasia, most likely in the area from Eastern Europe to South Asia. Because R1a1a is by far the most abundant form of R1a, all studies have focused on R1a1a. There are many proposals on the origin of R1a1a, including Eastern Europe, and Central, Western and Southern Asia. The most recent studies indicate that Asian areas are a slightly more likely point of origin than Europe.
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Phylogeny (Family Tree)
The most recent publications on this subject have increased knowledge of the complexity of R1a.[2][3] By far, the most commonly found type of R1a, now known as R1a1a, represents only one sub-branch of a bigger "family tree". Each major branching of this tree is identified by a corresponding set of known SNP mutations which are used to test individuals, and define the relationships between branches of descent. In particular, the discovery of the mutation M420, has defined a previously unidentified category of lineages, which are distant relatives of R1a1a.
Roots of R1a
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R1a evolved from a male-line ancestor who was in haplogroup R1. R1 is defined by SNP mutation M173. The R1a clade can be distinguished by several unique markers such as M420. R1a also has a similarly common sister-clade, called R1b, which also has M173, but is distinguished by its M343 marker. 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" implying that R1a's origin may perhaps be somewhere near these regions.
Different meanings of "R1a"
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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 clades or haplogroups. For example, the mutation called "M173" currently identifies R1. Thus R1 can also be called "R-M173". There are also "paragroups", which have an unknown number of branches. For example, men with R1* have the M173 marker, but they have no known defining mutations (e.g. neither M420 nor M343) other than those that identify R1. When a new branching in a tree is discovered, for example a branch within a paragroup, phylogenetic names need to change, while the marker-derived clade names remain the same.
The naming system commonly used for this haplogroup remains inconsistent in different published sources, and requires some explanation. A family tree for R1a was proposed in 2009 and appeared in published surveys in late 2009.[5][6] Prior to 2009 the mutation SRY1532.2 was commonly used to identify "R1a".[7] This is also how the term R1a is most often used in publications well into 2009. With the new phylogenetic tree, "R1a" is increasingly used to refer to a broader family. This includes the "old" R1a, but with the addition 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 R1a family tree as a whole can be divided into three major levels of branching, with a large number of currently defined sub-clades within the dominant and best known branch, R1a1a. The following summary is based upon the large survey of Underhill et al. (2009) as described in sections below and comparing the old and new naming systems.
R1a (R-M420)
This is the broadest definition of R1a, defined by the mutation M420. It is known to have at least two branches: R1a1 (see next section), which makes up the vast majority, and R1a*, the paragroup. In this newest definition, R1a* is defined as M420 positive but SRY1532.2 negative. However within the pre-2009 scheme, a minority of M420 positive cases would have been classified within R1* (R-M173*; see cladogram on right).
Underhill et al. (2009) found only isolated samples of the paragroup, 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 populations showed no sign of this category so far. Mutations understood to be equivalent to M420 include M449, M511, M513, L62, and L63.[2][8]
R1a1 (R-SRY1532.2)
R1a1 is currently defined by SRY1532.2, also referred to as SRY10831.2. This family of lineages is dominated by one very large and well-defined R1a1a branch, which is positive for M17 and M198 (see below). The paragroup R1a1* (old R1a*) is positive for the SRY1532.2 marker but lacks either the M17 or M198 markers.
Underhill et al. (2009) again found only limited examples of the R1a1* paragroup, looking at many different surveys. However it does appear to be spread over a wider geographical range in Eurasia than the R1a* paragroup 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.[2]
R1a1a (R-M17 or R-M198)
R1a1a (old R1a1) makes up the vast majority of all R1a, over its entire geographic range, and most statistical or other analysis of R1a is therefore by definition focused upon it. It is defined in various articles by SNP mutations M17 or M198, which always appears together in the same men. SNP mutations understood to be always occurring with M17 and M198 include M417, M512, M514, M515.[2] The vast majority of R1a1a has not yet been categorized into branches defined by mutations, and is therefore referred to as R1a1a*. However, this paragroup also has several known sub-clades of its own. So far, eight sub-clades of R1a1a are defined, however only a few are so far known to have significant frequencies.
R1a1a subclades
So far, 8 sub-clades of R1a1a are known, R1a1a1 to R1a1a8. Currently, only one of these subclades is known to have significant frequencies, R1a1a7. R1a1a7 is defined by M458 and was first described in Underhill et al. (2009). In Eurasia, M458 is found primarily Europe, with its highest frequencies in Central and Southern Poland. R1a1a7 has its own subclade, defined by the M334 marker.
| Region | People | N | R1a1a-M17 | R1a1a6-M434 | ||
| Number | Freq. (%) | Number | Freq. (%) | |||
| Pakistan | Baloch | 60 | 9 | 15% | 5 | 8% |
| Pakistan | Makrani | 60 | 15 | 25% | 4 | 7% |
| Middle East | Oman | 121 | 11 | 9% | 3 | 2.5% |
| Pakistan | Sindhi | 134 | 65 | 49% | 2 | 1% |
| Table only shows positive sets from N = 3667 derived from 60 Eurasian populations sample, Underhill et al. (2009) | ||||||
Concerning other sub-clades, R1a1a3 is defined by the M64.2, M87, and M204 SNPs and is apparently rare, for example found in 1 of 117 males typed in southern Iran.[9] And R1a1a6, defined by M434, was announced in Underhill et al. (2009). M434 was detected in 14 people (out of 3667 people tested) all in a restricted geographical range from Pakistan to Oman. This is likely to reflect a recent mutation that took place in the area of Pakistan.
Distribution of R1a1a (R-M17 or R-M198)
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 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.[10][11][12] 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.[10][13][14] R1a lines propogated north-eastward and are scattered amoung certain indigenous Eastern Siberians, including:Kamchatkans and Chukotkans, and peaking in Itel'man at 22%.[15]
South Asia
In South Asia high levels R1a 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[16] 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[17].
Middle East and Caucasus
R1a has been found in various forms, in most parts of 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.[18][19][20]
The uneven distribution in Iran has been the subject of special comment from Wells et al. (2001), who noted that in the western part of the country, Iranians appear to be more similar genetically to Semitic Speakers to the west, with low R1a levels while males of Eastern parts of iran carried 35% R1a. 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".
Turkey also shows high but unevenly distributed R1a levels amongst some sub-populations. For example Nasidze et al. (2005) found relatively high levels amongst Kurds (12%) and Zazas 26%.
Further to the north of these Middle Eastern regions on the other hand, R1a levels start to increase in the Caucasus, once again in an uneven way. Several populations studied have shown no sign of R1a, while highest levels so far discovered in the region appears to belong to speakers of the Karachay-Balkar language amongst whom about one quarter of men tested so far are in haplogroup R1a1a*.[2]
Europe
In Europe, R1a, again almost entirely in the R1a1a sub-clade, is found at highest levels among peoples of Eastern European descent (Sorbs, Poles, Russians and Ukranians; 50 to 65%).[21][22][23] Levels in Hungarians have been noted between 20 and 60% [24] 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.[23][25][26]. In the Baltic countries R1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).[27]
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.[28][29]
There is a significant presence in peoples of Scandinavian descent, with highest levels in Norway and Iceland, where between 20 and 30% of men are in R1a.[30][31] Vikings and Normans may have also carried the R1a lineage westward; accounting for at least part of the small presence in the British Isles.[32][33][34][35]
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
| STR site |
Frequency | |
| R1a1a(xM458) | R1a1a7 | |
| DYS19 | 16 | 16 |
| DYS388 | 12 | 12 |
| DYS389I | 13 | 13 |
| DYS389II | 17 | 16 |
| DYS390 | 25 | 25 |
| DYS391 | 11 | 10 |
| DYS392 | 11 | 11 |
| DYS393 | 13 | 13 |
| DYS439 | 10 | 11 |
| A7.2 | 10 | 10 |
Most discussions of R1a origins concern the dominant R1a1a (RM17 or R-M198) sub-clade. There are two foci of high frequency of R1a1a, one in South Asia, near North India, and the other in Eastern Europe, in the area of the Ukraine. Until 2009 claims regarding the oldest R1a bearing populations varied greatly between different articles, with Eastern Europe and South Asia being the main contenders. Such studies look at each region to see how different that R1a men are at specific unstable points on the Y chromosome known as STRs or microsatellites. Higher variance in a region is seen as an indicator that a haplogroup might have been present longer in that region.
In 2009, several large studies of both old and new STR data, including Mirabal et al. (2009), Underhill et al. (2009), and Klyosov (2009) concluded that not only are there are two separate "poles of the expansion" with similar ages, but also that of these two poles, Asian R1a1a is apparently older than European R1a1a. The data is therefore said to be more consistent with Asian origins for R1a1a, as opposed to European origins, with a particular focus remaining upon South Asia.[36]
Central Eurasian origin proposals
Cordaux et al. (2004) argued, citing data from 3 earlier publications, that R-M17 (R1a1a) Y chromosomes most probably have a central Asian origin.[37] 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.
Recently, looking at Chinese STR data not included in other studies Klyosov (2009) concluded that the common source of Indian and European R1a must be somewhere near the modern Chinese ethnic groups known as the Hui, Bolan, Dongxiang and Sala.
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 Indo-European 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 four 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.[38]
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, such as used by Klyosov (2009), 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. Klyosov (2009), on the other hand, would see this as a period for a very early branching within R1a1a, detectable from STR data rather than SNP data, wherein early European and Indian branches of R1a1a settled into their early positions. The initial homeland of European R1a1a in this scenario was on the very edge of Europe, near the Urals.
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".[39]
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.[40]
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.[41] 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.)[42]
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.[43][28][44] This was probably R1a1a* (M17/M198 positive, M458 negative) according to Underhill et al. (2009).
According to Klyosov (2009) however, there was a movement of R1a1a from Europe to India during this period, and it was associated with Indoeuropean language and culture. The author believes this flow originated on the edge of Europe, near the Urals. It should be noted that according to this scenario Indian R1a1a is made up of two components, one which came from the direction of Europe and one which arrived much earlier.
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
| Location | TD |
| W. India | 15,800 |
| Pakistan | 15,000 |
| Nepal | 14,200 |
| India | 14,000 |
| Oman | 12,500 |
| N. India | 12,400 |
| S. India | 12,400 |
| Caucasus | 12,200 |
| E. India | 11,800 |
| Poland | 11,300 |
| Slovakia | 11,200 |
| Crete | 11,200 |
| Germany | 9,900 |
| Denmark | 9,700 |
| UAE | 9,700 |
Several other studies suggest R1a lineages generally may have their origins in South Asia [16][45][46].
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). 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.[47] On the other hand, some authors have not accepted this association.[48]
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[49]). 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
- Genetic history of Europe
- Genetics and Archaeogenetics of South Asia
- Y-DNA haplogroups by ethnic groups
- Nordic R1a Y-DNA Project
- Somerled
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Human Y-chromosome DNA (Y-DNA) haplogroups (by ethnic groups · famous haplotypes) |
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Notes
- ^ Karafet et al. (2008)
- ^ a b c d e f g Underhill et al. (2009)
- ^ Underhill et al. (2009)
- ^ Identified by the authors with the standardized SNP reference rs34351054.
- ^ Underhill et al. (2009)
- ^ ISOGG phylogenetic tree
- ^ SRY1532.2 is also known as SRY10831.2
- ^ ISOGG phylogeny webpage 2009
- ^ Regueiro et al. (2006)
- ^ 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)
- ^ a b Sharma et al. (2007)
- ^ Kivisild et al. (2003)
- ^ Flores et al. (2005)
- ^ Nasidze et al. (2004)
- ^ Nasidze et al. (2005)
- ^ 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 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.
- ^ Rosser et al. (2000)
- ^ Pericic et al. (2005)
- ^ Kasperaviciūte et al. (2005)
- ^ a b Haak et al. (2008)
- ^ The Ysearch number for the Eulau remains is 2C46S.
- ^ Bowden et al. (2008)
- ^ Dupuy et al. (2005)
- ^ Irish Heritage DNA Project, R1 and R1a
- ^ Passarino et al. (2002)
- ^ Capelli et al. (2003)
- ^ Garvey, D. "Y Haplogroup R1a1". http://freepages.genealogy.rootsweb.com/%7Edgarvey/DNA/hg/YCC_R1a1.html. Retrieved 2007-04-23.
- ^ 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. Klyosov (2009) presents a more complex scenario in which R1a1a originated in South Siberia, branches headed to Europe and India, and then a branch from Europe also went to India.
- ^ Wells et al. (2001), Semino et al. (2000), and Quintana-Murci et al. (2001)
- ^ 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.
- ^ Dupuy et al. (2005)
- ^ see: Semino et al. (2000), Passarino et al. (2001), Passarino et al. (2002) and Wells (2002)
- ^ Keyser et al. (2009)
- ^ Lalueza-Fox et al. (2004)
- ^ Schilz (2006)
- ^ Bouakaze et al. (2007)
- ^ Sengupta et al. (2005)
- ^ Sahoo et al. (2006)
- ^ 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 [R1a1] than their Dravidian-speaking neighbours, the Yadhavas and Kallars, adding to the evidence that M17 [R1a1] is a diagnostic Indo-Iranian marker.
- ^ 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.
- ^ The authors also refer here to their references 14, Weale et al. (2001), and 41, Regueiro et al. (2006)]
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