|Possible time of origin||22,000 YBP (Sharma 2009)|
|Possible place of origin||Eurasia (see text).|
|Descendants||Haplogroup R1a-Z282 (Europe), R1a-Z93 (Asia)|
|Defining mutations||R1a: L62, L63, L120, M420, M449, M511, M513
R1a1a: M17, M198, M512, M514, M515, L168, L449, L457, L566
|Highest frequencies||See List of R1a frequency by population|
Haplogroup R1a, or haplogroup R-M420, is a Y DNA haplogroup which is distributed in a large region in Eurasia, extending from South Asia and southern Siberia to Central Europe and Scandinavia. The distribution of R1-Z282 (including R1-Z280) in Central- and Eastern Europe and R1-Z93 in Asia suggests that Ra diversified within the Eurasian Steppes or the Middle East and Caucasus region or "in the vicinity of present-day Iran."
The SNP mutation R-M420 was discovered after R-M17 (R1a1a1), which resulted in a reorganization of the lineage in particular establishing a new paragroup (designated R-M420*) for the relatively rare lineages which are not in the R-SRY10831.2 (R1a1) branch leading to R-M17.
- 1 Origins
- 2 Phylogeny
- 2.1 Topology
- 2.2 Haplogroup R
- 2.3 R-M173 (R1)
- 2.4 R-M420 (R1a)
- 2.5 R-SRY1532.2 (R1a1)
- 2.6 R-M17/M198 (R1a1a)
- 2.7 R1a1a1 (RM-417)
- 3 Geographic distribution of R1a1a
- 4 R1a and the Indo-Europeans
- 5 Popular science
- 6 Historic naming of "R1a"
- 7 See also
- 8 In art
- 9 Notes
- 10 References
- 11 Sources
- 12 Further reading
- 13 External links
The modern distribution of R1a1a has two widely separated areas of high frequency: South Asia and Eastern Europe. Some researchers have claimed that South Asian populations had the highest STR diversity within R1a1a. Other studies have proposed Eastern European, Central Asian and even West Asian origins for R1a1a.
According to Pamjav et al. (2012), "Inner and Central Asia is an overlap zone for the R1a1-Z280 and R1a1-Z93 lineages [which] implies that an early differentiation zone of R1a1-M198 conceivably occurred somewhere within the Eurasian Steppes or the Middle East and Caucasus region as they lie between South Asia and Central- and Eastern Europe." A large, 2014 study by Peter A Underhill et al., using 16,244 individuals from over 126 populations from across Eurasia, concluded there was compelling evidence, that "the initial episodes of haplogroup R1a diversification likely occurred in the vicinity of present-day Iran."
The R1a family tree now has three major levels of branching, with the largest number of defined subclades within the dominant and best known branch, R1a1a (which will be found with various names; in particular, as "R1a1" in relatively recent but not the latest literature.)
- R (R-M207)
- R1 (R-M173)
- R1a (M420) (Eastern Europe, Asia)
- R1a1 (M459/PF6235, SRY1532.2/SRY10831.2)
- R1a1 (M459)
- R1a1a (M17, M198)
- R1a1a1 (M417, page7)
- R1a1a1a (CTS7083/L664/S298)
- R1a1a1b (S224/Z645, S441/Z647)
- R1a1a1b1 (PF6217/S339/Z283)
- R1a1a1b1a (Z282) [R1a1a1a*] (Z282) (Eastern Europe)
- R1a1a1b1a1 [The old topological code is R1a1a1b*，which is outdated and might lead to some confusion.] (M458) [R1a1a1g] (M458)
- R1a1a1b1a2 (S466/Z280, S204/Z91)
- R1a1a1b1a3 (Z284) [R1a1a1a1] (Z284)
- R1a1a1b1a (Z282) [R1a1a1a*] (Z282) (Eastern Europe)
- R1a1a1b2 (F992/S202/Z93) [R1a1a2*] (Z93, M746)(Asia)
- R1a1a1b2a (F3105/S340/Z94, L342.2/S278.2) [R1a1b2a*] (Z95) R-Z94 (Z94/F3105/S340, Z95/F3568)
- R1a1a1b1 (PF6217/S339/Z283)
- [R1a1a1c] (M64.2, M87, M204)
- [R1a1a1d] (P98)
- [R1a1a1e] (PK5)
- R1a1a1 (M417, page7)
- R1b (M343) (Western Europe)
R1a, distinguished by several unique markers including the M420 mutation, is a subclade of Haplogroup R-M173 (previously called R1). R1a has the sister-subclades Haplogroup R1b-M343, and the paragroup R-M173*.
R-M420, defined by the mutation M420, has two branches: R-SRY1532.2, defined by the mutation SRY1532.2, which makes up the vast majority; and R-M420*, the paragroup, defined as M420 positive but SRY1532.2 negative. (In the 2002 scheme, this SRY1532.2 negative minority was one part of the relatively rare group classified as the paragroup R1*.) Mutations understood to be equivalent to M420 include M449, M511, M513, L62, and L63.
Only isolated samples of the new paragroup R-M420* were found by Underhill 2009, mostly in the Middle East and Caucasus: 1/121 Omanis, 2/150 Iranians, 1/164 in the United Arab Emirates, and 3/612 in Turkey. Testing of 7224 more males in 73 other Eurasian populations showed no sign of this category.
R1a1 is defined by SRY1532.2 or SRY10831.2), understood to always include SRY10831.2, M448, L122, M459, and M516.) This family of lineages is dominated by M17 and M198. In contrast, paragroup R-SRY1532.2* lacks either the M17 or M198 markers.
The R-SRY1532.2* paragroup is apparently less rare than R1*, but still relatively unusual, though it has been tested in more than one survey. Underhill et all. (2009) reported 1/51 in Norway, 3/305 in Sweden, 1/57 Greek Macedonians, 1/150 Iranians, 2/734 ethnic Armenians, and 1/141 Kabardians. Sahoo et al. (2006) reported R-SRY1532.2* for 1/15 Himachal Pradesh Rajput samples.
The following SNPs are associated with R1a1a:
|SNP||Mutation||Y-position (NCBI36)||Y-position (GRCh37)||RefSNP ID|
R1a1a1 (RM-417) is the most widely found subclade, in two variations which are found respectively in Europe (R1a1a1b1 (R-Z282) ([R1a1a1a*] (R-Z282) (Underhill 2014/2015)) and Central and South Asia (R1a1a1b2 (R-Z93) ([R1a1a2*] (R-Z93) Underhill 2014/2015)).
R1a1a1b1a (R-Z282) (Eastern Europe)
This large subclade appears to encompass most of the R1a1a found in Europe.
- R1a1a1b1a [R1a1a1a* (Underhill (2014))] (R-Z282*) occurs in northern Ukraine, Belarus, and Russia at a frequency of ~20%. (Underhill et al. 2014)
- R1a1a1b1a3 [R1a1a1a1 (Underhill (2014))] (R-Z284) occurs in Northwest Europe and peaks at ~20% in Norway. (Underhill et al. 2014)
- R1a1a1c (M64.2, M87, M204) is apparently rare: it was found in 1 of 117 males typed in southern Iran.
R-M458 is a mainly Slavic SNP, characterized by its own mutation, and was first called cluster N. Underhill et al. (2009) found it to be present in modern European populations roughly between the Rhine catchment and the Ural Mountains and traced it to "a founder effect that [...] falls into the early Holocene period, 7.9±2.6 KYA." M458 was found in one skeleton from a 14th-century grave field in Usedom, Mecklenburg-Vorpommern, Germany. The paper by Underhill et al. (2009) also reports a surprisingly high frequency of M458 in some Northern Caucasian populations (for example 27.5% among Karachays and 23.5% among Balkars, 7.8% among Karanogays and 3.4% among Abazas).
R1a1a1b1a1a (R-L260) (Gwozdz's cluster P)
R1a1a1b1a1a (R-L260), commonly referred to as West Slavic or Polish, is a subclade of the larger parent group R-M458, and was first identified as an STR cluster by Pawlowski 2002 and then by Gwozdz 2009. Thus, R-L260 was what Gwozdz 2009 called cluster "P." In 2010 it was verified to be a haplogroup identified by its own mutation (SNP). It apparently accounts for about 8% of Polish men, making it the most common subclade in Poland. Outside of Poland it is less common (Pawlowski 2002). In addition to Poland, it is mainly found in the Czech Republic and Slovakia, and is considered "clearly West Slavic." The founding ancestor of R-L260 is estimated to have lived between 2000 and 3000 years ago, i.e. during the Iron Age, with significant population expansion less than 1,500 years ago.
R-M334 ([R1a1a1g1], a subclade of [R1a1a1g] (M458) c.q. R1a1a1b1a1 (M458)) was found by Underhill et al. (2009) only in one Estonian man and may define a very recently founded and small clade.
R1a1a1b1a2 (S466/Z280, S204/Z91)
R1a1a1b1a2b3* (Gwozdz's Cluster K)
R1a1a1b1a2b3* (M417+, Z645+, Z283+, Z282+, Z280+, CTS1211+, CTS3402, Y33+, CTS3318+, Y2613+) (Gwozdz's Cluster K) is a STR based group that is R-M17(xM458). This cluster is common in Poland but not exclusive to Poland.
R1a1a1b2 (R-Z93) (Asia)
This large subclade appears to encompass most of the R1a1a found in Asia.
- R1a1a1b2 [R1a1a2* (Underhill (2014))] (R-Z93) is most common (>30%) in the South Siberian Altai region of Russia, cropping up in Kyrgyzstan (6%) and in all Iranian populations (1–8%).
- R1a1a1b2a* (R-Z2125): This subgroup occurs at highest frequencies in Kyrgyzstan and in Afghan Pashtuns (>40%). At a frequency of >10% it is also observed in other Afghan ethnic groups and in some populations in the Caucasus and Iran.
|Number||Freq. (%)||Number||Freq. (%)|
|Table only shows positive sets from N = 3667 derived from 60 Eurasian populations sample.|
- R1a1b2a1* (R-M560 is very rare and was only observed in four samples: two Burushaski speakers (north Pakistan), one Hazara (Afghanistan), and one Iranian Azerbaijani.
- R1a1b2a2* (R-M780) occurs at high frequency in South Asia: India, Pakistan, Afghanistan, and the Himalayas. The group also occurs at >3% in some Iranian populations and is present at >30% in Roma from Croatia and Hungary.
Geographic distribution of R1a1a
In Afghanistan, R1a1a is found at 51% among the Pashtuns who are the largest ethnic group in Afghanistan, 50% among the Kyrgyz, and 30% among the Tajiks. It is less frequent among the Hazaras (7%) and the Turkic-speaking Uzbeks (18%) (Haber 2012).
In India, high frequencies of this haplogroup is observed in West Bengal Brahmins (72%)(Sengupta 2005) to the east, Konkanastha Brahmins (48%) (Sengupta 2005) to the west, Khatris (67%)(Underhill 2009) in the north and Iyenger Brahmins (31%)(Sengupta 2005) in the south. It has also been found in several South Indian Dravidian-speaking Adivasis including the Chenchu (26%) and the Valmikis of Andhra Pradesh and the Kallar of Tamil Nadu suggesting that R1a1a is widespread in Tribal Southern Indians (Kivisild 2003).
Besides these, studies show high percentages in regionally diverse groups such as Manipuris (50%)(Underhill 2009) to the extreme North East and in Punjab (47%)(Kivisild 2003) to the extreme North West.
In Pakistan it is found at 71% among the Mohanna tribe in Sindh province to the south and 46% among the Baltis of Gilgit-Baltistan to the north (Underhill 2009). Among the Sinhalese of Sri Lanka, 23% were found to be R1a1a (R-SRY1532) positive in a sample size of 87 subjects. Hindus of Terai region of Nepal show it at 69% (Fornarino 2009).
The frequency of R1a1a is comparatively low among some Turkic-speaking groups including Turks, Azeris, Kazakhs, and Yakuts, yet levels are higher (19 to 28%) in certain Turkic or Mongolic-speaking groups of Northwestern China, such as the Bonan, Dongxiang, Salar, and Uyghurs.(Wells 2001, Wang 2003, and Zhou 2007)
R1a1a 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 Kuwait, Turkey and Iran. The Shimar (Shammar) Bedouin tribe in Kuwait show the highest frequency in the Middle East at 43%.(Mohammad 2009,Nasidze 2004, and Nasidze 2005)
Wells 2001, noted that in the western part of the country, Iranians show low R1a1a levels, while males of eastern parts of Iran carried up to 35% R1a1a. Nasidze 2004 found R1a1a in approximately 20% of Iranian males from the cities of Tehran and Isfahan. Regueiro 2006 in a study of Iran, noted much higher frequencies in the south than the north.
A newer Study has found 20.3% R-M17* among Kurdish samples which were taken in the Kurdistan Province in western Iran, 9.7% among Mazandaranis in North Iran in the province of Mazandaran, 9.4% among Gilaks in province of Gilan, 12.8% among Persian and 17.6% among Zoroastrians in Yazd, 18.2% among Persians in Isfahan, 20.3% among Persians in Khorasan, 16.7% Afro-Iranians, 18.4% Qeshmi "Gheshmi", 21.4% among Persian Speaking Bandari people in Hormozgan and 25% among the Baloch people in Sistan and Baluchestan Province (Grugni 2012).
Further to the north of these Middle Eastern regions on the other hand, R1a1a levels start to increase in the Caucasus, once again in an uneven way. Several populations studied have shown no sign of R1a1a, while highest levels so far discovered in the region appears to belong to speakers of the Karachay-Balkar language among whom about one quarter of men tested so far are in haplogroup R1a1a (Underhill 2009).
In Europe, the R1a1 sub-clade, is found at highest levels among peoples of Eastern European descent (Sorbs, Poles, Russians and Ukrainians; 50 to 65%) (Balanovsky 2008, Behar 2003, and Semino 2000). In the Baltic countries R1a1a frequencies decrease from Lithuania (45%) to Estonia (around 30%) (Kasperaviciūte 2005). Levels in Hungarians have been noted between 20 and 60% (Battaglia 2008, Rosser 2000, Semino 2000, and Tambets 2004).
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 R1a1a (Bowden 2008 and (Dupuy 2005)). Vikings and Normans may have also carried the R1a1a lineage westward; accounting for at least part of the small presence in the British Isles (Passarino 2002 and Capelli 2003). In East Germany, where Haplogroup R1a1a reaches a peak frequency in Rostock at a percentage of 31.3%, it averages between 20%-30% (Kayser 2005).
Haplogroup R1a1a was found at elevated levels among a sample of the Israeli population who self-designated themselves as Levites and Ashkenazi Jews (Levites comprise approximately 4% of Jews). Behar reported R1a1a to be the dominant haplogroup in Ashkenazi Levites (52%), although rare in Ashkenazi Cohanim (1.3%). (Behar 2003).
In Southern Europe R1a1a is not common, but significant levels have been found in pockets, such as in the Pas Valley in Northern Spain, areas of Venice, and Calabria in Italy (Scozzari 2001). 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 (Pericić 2005, Rosser 2000, and Semino 2000).
The remains of a father and his two sons, from an archaeological site discovered in 2005 near Eulau (in Saxony-Anhalt, Germany) and dated to about 2600 BCE, tested positive for the Y-SNP marker SRY10831.2. The Ysearch number for the Eulau remains is 2C46S. The ancestral clade was thus present in Europe at least 4600 years ago, in association with one site of the widespread Corded Ware culture (Haak 2008).
R1a and the Indo-Europeans
The question of the origins of R1a1a is relevant to the ongoing debate concerning the urheimat of the proto-Indo-European people, and may also be relevant to the origins of the Indus Valley Civilisation. R1a shows a strong correlation with Indo-European languages of western Asia and eastern Europe, being most prevalent in Poland, Russia, and Ukraine and also observed in Pakistan, India and central Asia. The connection between Y-DNA R-M17 and the spread of Indo-European languages was first noted by T. Zerjal and colleagues in 1999. Ornella Semino and colleagues proposed a postglacial spread of the R1a1 gene during the Late Glacial Maximum, subsequently magnified by the expansion of the Kurgan culture into Europe and eastward. Spencer Wells suggests that the distribution and age of R1a1 points to an ancient migration corresponding to the spread by the Kurgan people in their expansion from the Eurasian steppe.
Haplogroup R1a has been found in the remains of people of the Corded Ware culture and Urnfield culture;[unreliable source?] as well as the burial of the remains of the Andronovo culture, the Pazyryk culture, Tagar culture and Tashtyk culture, the inhabitants of ancient Tanais, in the Tarim mummies, the aristocracy Xiongnu.
According to Underhill et al. (2014/2015) the diversification of Z93 and the "early urbanization within the Indus Valley also occurred at [5,600 years ago] and the geographic distribution of R1a-M780 (Figure 3d) may reflect this." Poznik et al. (2016) note that 'striking expansions' occurred within R1a-Z93 at ~4,500-4,000 years ago, which "predates by a few centuries the collapse of the Indus Valley Civilisation."
Mascarenhas et al. (2015) note that the expansion of Z93 from Transcaucasia into South Asia is compatible with "the archeological records of eastward expansion of West Asian populations in the 4th millennium BCE culminating in the so-called Kura-Araxes migrations in the post-Uruk IV period."
According to Lazaridis et al. (2016), "farmers related to those from Iran spread northward into the Eurasian steppe; and people related to both the early farmers of Iran and to the pastoralists of the Eurasian steppe spread eastward into South Asia." They further note that ANI "can be modelled as a mix of ancestry related to both early farmers of western Iran and to people of the Bronze Age Eurasian steppe."[note 1]
Bryan Sykes in his book Blood of the Isles gives imaginative names to the founders or "clan patriarchs" of major British Y haplogroups, much as he did for mitochondrial haplogroups in his work The Seven Daughters of Eve. He named R1a1a in Europe the "clan" of a "patriarch" Sigurd, reflecting the theory that R1a1a in the British Isles has Norse origins.
Historic naming of "R1a"
The historic naming system commonly used for R1a was inconsistent in different published sources, because it changed often; this requires some explanation.
In 2002, the Y Chromosome Consortium (YCC) proposed a new naming system for haplogroups (YCC 2002), which has now become standard. In this system, names with the format "R1" and "R1a" are "phylogenetic" names, aimed at marking positions in a family tree. Names of SNP mutations can also be used to name clades or haplogroups. For example, as M173 is currently the defining mutation of R1, R1 is also R-M173, a "mutational" clade name. When a new branching in a tree is discovered, some phylogenetic names will change, but by definition all mutational names will remain the same.
The widely occurring haplogroup defined by mutation M17 was known by various names, such as "Eu19", as used in (Semino 2000) in the older naming systems. The 2002 YCC proposal assigned the name R1a to the haplogroup defined by mutation SRY1532.2. This included Eu19 (i.e. R-M17) as a subclade, so Eu19 was named R1a1. Note, SRY1532.2 is also known as SRY10831.2 The discovery of M420 in 2009 has caused a reassignment of these phylogenetic names.(Underhill 2009 and ISOGG 2012) R1a is now defined by the M420 mutation: in this updated tree, the subclade defined by SRY1532.2 has moved from R1a to R1a1, and Eu19 (R-M17) from R1a1 to R1a1a.
More recent updates recorded at the ISOGG reference webpage involve branches of R-M17, including one major branch, R-M417.
|2002 Scheme proposed in (YCC 2002)||2009 Scheme as per (2009)||Latest ISOGG tree as per January 2011|
Y-DNA R-M207 subclades
Y-DNA backbone tree
|Evolutionary tree of human Y-chromosome DNA haplogroups [χ 1][χ 2]|
|A00||A0-T [χ 3]|
|A0||A1 [χ 4]|
|I||J||LT [χ 5]||K2|
|L||T [χ 6]||NO [χ 7]||K2b [χ 8]||K2c||K2d||K2e [χ 9]|
|N||O||K2b1 [χ 10]||P|
|K2b1a [χ 11]||K2b1b||K2b1c||M||P1||P2|
|K2b1a1||K2b1a2||K2b1a3||S [χ 12]||Q||R|
Artem Lukichev created an animation based on the Bashkir epic about the Ural, which outlined the history of the clusters of haplogroup R1: R1a and R1b.
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