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Haplogroup R1a

January 17, 2023

Haplogroup R1a, or haplogroup R-M420, is a human Y-chromosome DNA haplogroup which is distributed in a large region in Eurasia, extending from Scandinavia and Central Europe to southern Siberia and South Asia.[3][2]

Haplogroup R1a
Possible time of origin22,000 YBP[1] to 25,000[2] years ago
Possible place of originEastern Europe, South Asia, Central Asia, or West Asia
AncestorHaplogroup R1
DescendantsR1a-Z282, R1a-Z93
Defining mutations
  • R1a: L62, L63, L120, M420, M449, M511, M513
  • R1a1a: M17, M198, M512, M514, M515, L168, L449, L457, L566
Highest frequenciesSee List of R1a frequency by population
Map showing frequency of R1a haplogroup in Europe

While R1a originated c. 22,000[1] to 25,000[2] years ago, its subclade M417 (R1a1a1) diversified c. 5,800 years ago.[4] The place of origin of the subclade plays a role in the debate about the origins of Proto-Indo-Europeans.

The SNP mutation R-M420 was discovered after R-M17 (R1a1a), 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.

Origins

R1a origins

The split of R1a (M420) is computed to c. 22,000[1] or 25,000[2] years ago, which is the time of the last glacial maximum. A 2014 study by Peter A. Underhill et al., using 16,244 individuals from over 126 populations from across Eurasia, concluded that there was "a compelling case for the Middle East, possibly near present-day Iran, as the geographic origin of hg R1a".[2] The ancient DNA record has shown the first R1a during the Mesolithic in Eastern Hunter-Gatherers (from Eastern Europe),[5][6] and the earliest case of R* among Upper Paleolithic Ancient North Eurasians,[7] from which the Eastern Hunter-Gatherers predominantly derive their ancestry. [8]

Diversification of R1a1a1 (M417) and ancient migrations

 
R1a origins (Underhill 2009;[3] R1a1a origins (Pamjav et al. 2012); possible migration R1a to Baltic coast; and R1a1a oldest expansion and highest frequency (Underhill et al. 2014)

According to Underhill et al. (2014), the downstream R1a-M417 subclade diversified into Z282 and Z93 circa 5,800 years ago "in the vicinity of Iran and Eastern Turkey".[4][note 1] Even though R1a occurs as a Y-chromosome haplogroup among various languages such as Slavic and Indo-Iranian, 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 civilization. R1a shows a strong correlation with Indo-European languages of Southern and Western Asia, Central and Eastern Europe and to some extent Scandinavia[10][3] being most prevalent in Eastern Europe, West Asia, and South Asia. In Europe, Z282 is prevalent particularly while in Asia Z93 dominates. The connection between Y-DNA R-M17 and the spread of Indo-European languages was first noted by T. Zerjal and colleagues in 1999.[11]

Proposed steppe dispersal of R1a1a

See also: Indo-European migrations § Proto-Indo-Europeans, and Indo-Aryan migration § Haplogroup R1a and related haplogroups

Semino et al. (2000) proposed Ukrainian origins, and a postglacial spread of the R1a1 haplogroup during the Late Glacial Maximum, subsequently magnified by the expansion of the Kurgan culture into Europe and eastward.[12] Spencer Wells proposes Central Asian origins, suggesting 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.[13] According to Pamjav et al. (2012), R1a1a diversified in the Eurasian Steppes or the Middle East and Caucasus region:

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.[14]

Three genetic studies in 2015 gave support to the Kurgan theory of Gimbutas regarding the Indo-European Urheimat. According to those studies, haplogroups R1b and R1a, now the most common in Europe (R1a is also common in South Asia) would have expanded from the Pontic–Caspian steppes, along with the Indo-European languages; they also detected an autosomal component present in modern Europeans which was not present in Neolithic Europeans, which would have been introduced with paternal lineages R1b and R1a, as well as Indo-European languages.[15][16][17]

Silva et al. (2017) noted that R1a in South Asia most "likely spread from a single Central Asian source pool, there do seem to be at least three and probably more R1a founder clades within the Subcontinent, consistent with multiple waves of arrival."[18] According to Martin P. Richards, co-author of Silva et al. (2017), the prevalence of R1a in India was "very powerful evidence for a substantial Bronze Age migration from central Asia that most likely brought Indo-European speakers to India."[19][note 2]

Proposed South Asian origins

Kivisild et al. (2003) have proposed either South or West Asia,[20][note 3] while Mirabal et al. (2009) see support for both South and Central Asia.[10] Sharma et al.(2009) showcased the existence of R1a in India beyond 18,000 years to possibly 44,000 years in origin.[1]

South Asian populations have the highest STR diversity within R1a1a,[21][22][10][3][1][23] and subsequent older TMRCA datings,[citation needed] and R1a1a is present among both higher (Brahmin) castes and lower castes, although the frequency is higher among Brahmin castes. Nevertheless, the oldest TMRCA datings of the R1a haplogroup occur in the Saharia tribe, a scheduled caste of the Bundelkhand region.[1][23] From these findings some researchers have concluded that R1a1a originated in South Asia,[22][1][note 4][note 5] excluding a more recent, yet minor, genetic influx from Indo-European migrants in northwestern regions such as Afghanistan, Balochistan, Punjab, and Kashmir.[22][21][3]

However, this diversity, and the subsequent older TMRCA-datings, can also be explained by the historically high population numbers,[note 6] which increases the likelihood of diversification and microsatellite variation.[19][18] According to Sengupta et al. (2006), "[R1a1 and R2] could have actually arrived in southern India from a southwestern Asian source region multiple times."[21][note 7] However, Sengupta also described in this article:

We found that the influence of Central Asia on the pre-existing gene pool was minor. The ages of accumulated microsatellite variation in the majority of Indian haplogroups exceed 10,000–15,000 years, which attests to the antiquity of regional differentiation. Therefore, our data do not support models that invoke a pronounced recent genetic input from Central Asia to explain the observed genetic variation in South Asia.

This suggests that the origins of paternal haplogroup R1a point to the Indian subcontinent and not Central Asia.

Proposed Yamnaya origins

 
European middle-Neolithic period. Comb Ware culture c. 4200 – c. 2000 BCE
 
Corded Ware culture (c. 2900 – c. 2350 BCE

David Anthony considers the Yamnaya culture to be the Indo-European Urheimat.[25][26] According to Haak et al. (2015), a massive migration from the Yamnaya culture northwards took place c. 2,500 BCE, accounting for 75% of the genetic ancestry of the Corded Ware culture, noting that R1a and R1b may have "spread into Europe from the East after 3,000 BCE".[27] Yet, all their seven Yamnaya samples belonged to the R1b-M269 subclade,[27] but no R1a1a has been found in their Yamnaya samples. This raises the question where the R1a1a in the Corded Ware culture came from, if it was not from the Yamnaya culture.[28]

According to Marc Haber, the absence of haplogroup R1a-M458 in Afghanistan does not support a Pontic-Caspian steppe origin for the R1a lineages in modern Central Asian populations.[29] According to Leo Klejn, the absence of haplogroup R1a in Yamnaya remains (despite its presence in Eneolithic Samara and Eastern Hunter Gatherer populations) makes it unlikely that Europeans inherited haplogroup R1a from Yamnaya.[30] Archaeologist Barry Cunliffe has said that the absence of haplogroup R1a in Yamnaya specimens is a major weakness in Haak's proposal that R1a has a Yamnaya origin.[31]

Semenov & Bulat (2016) do argue for a Yamnaya origin of R1a1a in the Corded Ware culture, noting that several publications point to the presence of R1a1 in the Comb Ware culture.[32][note 8]

Proposed Transcaucasia and West Asian origins and possible influence on Indus Valley Civilization

See also: Kura–Araxes culture, Uruk period, and Indus Valley Civilisation § Possible near-Eastern Dravidian origins

Haak et al. (2015) found that part of the Yamnaya ancestry derived from the Middle East and that neolithic techniques probably arrived at the Yamnaya culture from the Balkans.[note 9] The Rössen culture (4,600–4,300 BC), which was situated on Germany and predates the Corded Ware culture, an old subclade of R1a, namely L664, can still be found.[note 10]

Part of the South Asian genetic ancestry derives from west Eurasian populations, and some researchers have implied that Z93 may have come to India via Iran[34] and expanded there during the Indus Valley civilization.[2][35]

Mascarenhas et al. (2015) proposed that the roots of Z93 lie in West Asia, and proposed that "Z93 and L342.2 expanded in a southeasterly direction from Transcaucasia into South Asia",[34] noting that such an expansion 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."[34] Yet, Lazaridis noted that sample I1635 of Lazaridis et al. (2016), their Armenian Kura-Araxes sample, carried Y-haplogroup R1b1-M415(xM269)[note 11] (also called R1b1a1b-CTS3187).[36][unreliable source?]

According to Underhill et al. (2014) the diversification of Z93 and the "early urbanization within the Indus Valley ... occurred at [5,600 years ago] and the geographic distribution of R1a-M780 (Figure 3d[note 12]) may reflect this."[2][note 13] Poznik et al. (2016) note that "striking expansions" occurred within R1a-Z93 at c. 4,500–4,000 years ago, which "predates by a few centuries the collapse of the Indus Valley Civilisation."[35][note 14]

However, according to Narasimhan et al. (2018), steppe pastoralists are a likely source for R1a in India.[38][note 15]

Phylogeny

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 such as "R1a1" in relatively recent but not the latest literature).

Topology

The topology of R1a is as follows (codes [in brackets] non-isogg codes):[9][39][verification needed][40][2][41] Tatiana et al. (2014) "rapid diversification process of K-M526 likely occurred in Southeast Asia, with subsequent westward expansions of the ancestors of haplogroups R and Q."[42]

  • P P295/PF5866/S8 (also known as K2b2).
  • R (R-M207)[40][9]
    • R*
    • R1 (R-M173)
      • R1*[40]
      • R1a (M420)[40] (Eastern Europe, Asia)[2]
        • R1a*[9]
        • R1a1[40] (M459/PF6235,[40] SRY1532.2/SRY10831.2[40])
          • R1a1 (M459)[40][9]
          • R1a1a (M17, M198)[40]
            • R1a1a1 (M417, page7)[40]
              • R1a1a1a (CTS7083/L664/S298)[40]
              • R1a1a1b (S224/Z645, S441/Z647)[40]
                • R1a1a1b1 (PF6217/S339/Z283)[40]
                  • R1a1a1b1a (Z282)[40] [R1a1a1a*] (Z282) [43]
                    • R1a1a1b1a1[40] [The old topological code is R1a1a1b*，which is outdated and might lead to some confusion.][43] (M458)[40][43] [R1a1a1g] (M458)[41]
                    • R1a1a1b1a2[40] (S466/Z280, S204/Z91)[40]
                      • R1a1a1b1a2a[40]
                      • R1a1a1b1a2b (CTS1211)[40] [R1a1a1c*] (M558)[43] [R-CTS1211] (V2803/CTS3607/S3363/M558, CTS1211/S3357, Y34/FGC36457)[9]
                        • R1a1a1b1a2b3* (M417+, Z645+, Z283+, Z282+, Z280+, CTS1211+, CTS3402, Y33+, CTS3318+, Y2613+) (Gwozdz's Cluster K)[39][verification needed]
                        • R1a1a1b1a2b3a (L365/S468)[40]
                    • R1a1a1b1a3 (Z284)[40] [R1a1a1a1] (Z284)[43]
                • R1a1a1b2 (F992/S202/Z93)[40] [R1a1a2*] (Z93, M746)[43]
                  • R1a1a1b2a (F3105/S340/Z94, L342.2/S278.2)[40] [R1a1b2a*] (Z95)[43] R-Z94 (Z94/F3105/S340, Z95/F3568)[9]
                    • R-Z2124 (Z2121/S3410, Z2124)[9]
                      • [R1a1b2a*] (Z2125)[43]
                        • [R1a1b2a*] (M434)[43] [R1a1a1f] (M434)[41]
                        • [R1a1b2a*] (M204)[43]
                    • [R1a1b2a1*] (M560)[43]
                    • [R1a1b2a2*] (M780, L657)[43] (India)[2]
                    • [R1a1b2a3*] (Z2122, M582)[43]
              • [R1a1a1c] (M64.2, M87, M204)[41]
              • [R1a1a1d] (P98)[41]
              • [R1a1a1e] (PK5)[41]
      • R1b (M343) (Western Europe)
    • R2 (India)

Haplogroup R

Haplogroup R phylogeny
 
R  (M207)   
 R1   (M173)   
  M420 

 R1a

  M343 

 R1b

 M173(xM420, M343) 

 R1*

R2 (M479)    

R* M207(xM173, M479)

R-M173 (R1)

R1a is distinguished by several unique markers, including the M420 mutation. It 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 (R1a)

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.[3][45]

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.[3]

This paragroup is now known as R1a2 (R-YP4141). It then has two branches R1a2a (R-YP5018) and R1a2b (R-YP4132).

R-SRY1532.2 (R1a1)

R1a1 is defined by SRY1532.2 or SRY10831.2 (understood to always include SRY10831.2, M448, L122, M459, and M516[3][46]). 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 al. (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.[3] Sahoo et al. (2006) reported R-SRY1532.2* for 1/15 Himachal Pradesh Rajput samples.[22]

R-M17/M198 (R1a1a)

The following SNPs are associated with R1a1a:

SNP Mutation Y-position (NCBI36) Y-position (GRCh37) RefSNP ID
M17 INS G 20192556 21733168 rs3908
M198 C->T 13540146 15030752 rs2020857
M512 C->T 14824547 16315153 rs17222146
M514 C->T 17884688 19375294 rs17315926
M515 T->A 12564623 14054623 rs17221601
L168 A->G 14711571 16202177 -
L449 C->T 21376144 22966756 -
L457 G->A 14946266 16436872 rs113195541
L566 C->T - - -

R-M417 (R1a1a1)

R1a1a1 (R-M417) is the most widely found subclade, in two variations which are found respectively in Europe (R1a1a1b1 (R-Z282) ([R1a1a1a*] (R-Z282) (Underhill 2014)[2]) and Central and South Asia (R1a1a1b2 (R-Z93) ([R1a1a2*] (R-Z93) Underhill 2014)[2]).

R-Z282 (R1a1a1b1a) (Eastern Europe)

This large subclade appears to encompass most of the R1a1a found in Europe.[14]

  • R1a1a1b1a [R1a1a1a* (Underhill (2014))] (R-Z282*) occurs in northern Ukraine, Belarus, and Russia at a frequency of c. 20%.[2]
  • R1a1a1b1a3 [R1a1a1a1 (Underhill (2014))] (R-Z284) occurs in Northwest Europe and peaks at c. 20% in Norway.[2]
  • R1a1a1c (M64.2, M87, M204) is apparently rare: it was found in 1 of 117 males typed in southern Iran.[47]
R-M458 (R1a1a1b1a1)
 
Frequency distribution of R-M458

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."[3] M458 was found in one skeleton from a 14th-century grave field in Usedom, Mecklenburg-Vorpommern, Germany.[48] 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).

R-L260 (R1a1a1b1a1a)

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 et al. 2002. In 2010 it was verified to be a haplogroup identified by its own mutation (SNP).[49] It apparently accounts for about 8% of Polish men, making it the most common subclade in Poland. Outside of Poland it is less common.[50] 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.[51]

R-M334

R-M334 ([R1a1a1g1],[41] a subclade of [R1a1a1g] (M458)[41] c.q. R1a1a1b1a1 (M458)[40]) was found by Underhill et al. (2009) only in one Estonian man and may define a very recently founded and small clade.[3]

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)[39][verification needed] is a STR based group that is R-M17(xM458). This cluster is common in Poland but not exclusive to Poland.[51]

R1a1a1b1a2b3a (R-L365)

R1a1a1b1a2b3a (R-L365)[40] was early called Cluster G.[citation needed]

R1a1a1b2 (R-Z93) (Asia)

Relative frequency of R-M434 to R-M17
Region People N R-M17 R-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 1.5%
Table only shows positive sets from N = 3667 derived from 60 Eurasian populations sample.[3]

This large subclade appears to encompass most of the R1a1a found in Asia, being related to Indo-European migrations (including Scythians, Indo-Aryan migrations and so on).[14]

  • R-Z93* or R1a1a1b2* (R1a1a2* in Underhill (2014)) is most common (>30%) in the South Siberian Altai region of Russia, cropping up in Kyrgyzstan (6%) and in all Iranian populations (1-8%).[2]
  • R-Z2125 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.[2]
    • R-M434 is a subclade of Z2125. It was detected in 14 people (out of 3667 people tested), all in a restricted geographical range from Pakistan to Oman. This likely reflects a recent mutation event in Pakistan.[3]
  • R-M560 is very rare and was only observed in four samples: two Burushaski speakers (north Pakistan), one Hazara (Afghanistan), and one Iranian Azerbaijani.[2]
  • 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.[2]

Geographic distribution of R1a1a

 
Distribution of R1a (purple) and R1b (red)

Pre-Historical

In Mesolithic Europe, R1a is characteristic of Eastern Hunter-Gatherers (EHGs).[52] A male EHG of the Veretye culture buried at Peschanitsa near Lake Lacha in Arkhangelsk Oblast, Russia c. 10,700 BCE was found to be a carrier of the paternal haplogroup R1a5-YP1301 and the maternal haplogroup U4a.[53][54][52] A Mesolithic male from Karelia c. 8,800 BCE to 7950 BCE has been found to be carrying haplogroup R1a.[55] A Mesolithic male buried at Deriivka c. 7000 BCE to 6700 BCE carried the paternal haplogroup R1a and the maternal U5a2a.[17] Another male from Karelia from c. 5,500 to 5,000 BC, who was considered an EHG, carried haplogroup R1a.[15] A male from the Comb Ceramic culture in Kudruküla c. 5,900 BCE to 3,800 BCE has been determined to be a carrier of R1a and the maternal U2e1.[56] According to archaeologist David Anthony, the paternal R1a-Z93 was found at Alexandria, Ukraine c. 4000 BCE, Sredny Stog culture, "the earliest known sample to show the genetic adaptation to lactase persistence (13910-T)."[57] R1a has been found in the Corded Ware culture,[58][59] in which it is predominant.[60] Examined males of the Bronze Age Fatyanovo culture belong entirely to R1a, specifically subclade R1a-Z93.[52][53][61]

Haplogroup R1a has later been found in ancient fossils associated with the Urnfield culture;[62] as well as the burial of the remains of the Sintashta,[16] Andronovo,[63] the Pazyryk,[64] Tagar,[63] Tashtyk,[63] and Srubnaya cultures, the inhabitants of ancient Tanais,[65] in the Tarim mummies,[66] and the aristocracy of Xiongnu.[67] The skeletal 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.[58]

Historical to recent Europe

In Europe, the R1a1 sub-clade is found at highest levels among peoples of Central and Eastern European descent, with results ranging from 35% to 65% among Czechs, Hungarians, Poles, Slovaks, western Ukrainians (particularly Rusyns), Belarusians, Moldovans, and Russians.[68][69][12] In the Baltics, R1a1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).[70][71][72][12][73]

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.[74][75] Vikings and Normans may have also carried the R1a1a lineage further out, accounting for at least part of the small presence in the British Isles, the Canary Islands, and Sicily.[76][77] In East Germany, Haplogroup R1a1a averages between 20 and 30%, with a peak in Rostock at 31.3%, it.[78]

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.[79][better source needed] The Balkans shows wide variation between areas with significant levels of R1a1a, for example 36–39% in Slovenia,[80] 27–34% in Croatia,[71][81][82][83][84] and over 30% in Greek Macedonia, but less than 10% in Albania, Kosovo and parts of Greece south of Olympus gorge.[85][72][12]

R1a is virtually composed only of the Z284 subclade in Scandinavia. In Slovenia, the main subclade is Z282 (Z280 and M458), although the Z284 subclade was found in one sample of a Slovenian. There is a negligible representation of Z93 in each region other than Turkey.[2] West Slavs and Hungarians are characterized by a high frequency of the subclade M458 and a low Z92, a subclade of Z280. Hundreds of Slovenian samples and Czechs lack the Z92 subclade of Z280, while Poles, Slovaks, Croats and Hungarians only show a very low frequency of Z92.[2] The Balts, East Slavs, Serbs, Macedonians, Bulgarians and Romanians demonstrate a ratio Z280>M458 and a high, up to a prevailing share of Z92.[2] Balts and East Slavs have the same subclades and similar frequencies in a more detailed phylogeny of the subclades.[86][87] The Russian geneticist Oleg Balanovsky speculated that there is a predominance of the assimilated pre-Slavic substrate in the genetics of East and West Slavic populations, according to him the common genetic structure which contrasts East Slavs and Balts from other populations may suggest the explanation that the pre-Slavic substrate of the East Slavs consisted most significantly of Baltic-speakers, which at one point predated the Slavs in the cultures of the Eurasian steppe according to archaeological and toponymic references.[note 16]

Asia

Central Asia

Zerjal et al. (2002) found R1a1a in 64% of a sample of the Tajiks of Tajikistan and 63% of a sample of the Kyrgyz of Kyrgyzstan.[88]

Haber et al. (2012) found R1a1a-M17 in 26.0% (53/204) of a set of samples from Afghanistan, including 60% (3/5) of a sample of Nuristanis, 51.0% (25/49) of a sample of Pashtuns, 30.4% (17/56) of a sample of Tajiks, 17.6% (3/17) of a sample of Uzbeks, 6.7% (4/60) of a sample of Hazaras, and in the only sampled Turkmen individual.[89]

Di Cristofaro et al. (2013) found R1a1a-M198/M17 in 56.3% (49/87) of a pair of samples of Pashtuns from Afghanistan (including 20/34 or 58.8% of a sample of Pashtuns from Baghlan and 29/53 or 54.7% of a sample of Pashtuns from Kunduz), 29.1% (37/127) of a pool of samples of Uzbeks from Afghanistan (including 28/94 or 29.8% of a sample of Uzbeks from Jawzjan, 8/28 or 28.6% of a sample of Uzbeks from Sar-e Pol, and 1/5 or 20% of a sample of Uzbeks from Balkh), 27.5% (39/142) of a pool of samples of Tajiks from Afghanistan (including 22/54 or 40.7% of a sample of Tajiks from Balkh, 9/35 or 25.7% of a sample of Tajiks from Takhar, 4/16 or 25.0% of a sample of Tajiks from Samangan, and 4/37 or 10.8% of a sample of Tajiks from Badakhshan), 16.2% (12/74) of a sample of Turkmens from Jawzjan, and 9.1% (7/77) of a pair of samples of Hazara from Afghanistan (including 7/69 or 10.1% of a sample of Hazara from Bamiyan and 0/8 or 0% of a sample of Hazara from Balkh).[90]

Malyarchuk et al. (2013) found R1a1-SRY10831.2 in 30.0% (12/40) of a sample of Tajiks from Tajikistan.[91]

Ashirbekov et al. (2017) found R1a-M198 in 6.03% (78/1294) of a set of samples of Kazakhs from Kazakhstan. R1a-M198 was observed with greater than average frequency in the study's samples of the following Kazakh tribes: 13/41 = 31.7% of a sample of Suan, 8/29 = 27.6% of a sample of Oshaqty, 6/30 = 20.0% of a sample of Qozha, 4/29 = 13.8% of a sample of Qypshaq, 1/8 = 12.5% of a sample of Tore, 9/86 = 10.5% of a sample of Jetyru, 4/50 = 8.0% of a sample of Argyn, 1/13 = 7.7% of a sample of Shanyshqyly, 8/122 = 6.6% of a sample of Alimuly, 3/46 = 6.5% of a sample of Alban. R1a-M198 also was observed in 5/42 = 11.9% of a sample of Kazakhs of unreported tribal affiliation.[92]

South Asia

In South Asia, R1a1a has often been observed in a number of demographic groups.[22][21]

In India, high frequencies of this haplogroup is observed in West Bengal Brahmins (72%)[21] to the east, Gujarat Lohanas (60%) [3] to the west, Khatris (67%)[3] in the north and Iyengar Brahmins (31%)[21] 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, Kota (22.58%)[93] and the Kallar of Tamil Nadu suggesting that R1a1a is widespread in Tribal Southern Indians.[20]

Besides these, studies show high percentages in regionally diverse groups such as Manipuris (50%)[3] to the extreme North East and among Punjabis (47%)[20] 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.[3] Among the Sinhalese of Sri Lanka, 23% were found to be R1a1a (R-SRY1532) positive.[94] Hindus of Chitwan District in the Terai region Nepal show it at 69%.[95]

East Asia

The frequency of R1a1a is comparatively low among some Turkic-speaking groups like 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.[13][96][97]

A Chinese paper published in 2018 found R1a-Z94 in 38.5% (15/39) of a sample of Keriyalik Uyghurs from Darya Boyi / Darya Boye Village, Yutian County, Xinjiang (于田县达里雅布依乡), R1a-Z93 in 28.9% (22/76) of a sample of Dolan Uyghurs from Horiqol township, Awat County, Xinjiang (阿瓦提县乌鲁却勒镇), and R1a-Z93 in 6.3% (4/64) of a sample of Loplik Uyghurs from Karquga / Qarchugha Village, Yuli County, Xinjiang (尉犁县喀尔曲尕乡). R1a(xZ93) was observed only in one of 76 Dolan Uyghurs.[98] Note that Darya Boyi Village is located in a remote oasis formed by the Keriya River in the Taklamakan Desert. A 2011 Y-DNA study found Y-dna R1a1 in 10% of a sample of southern Hui people from Yunnan, 1.6% of a sample of Tibetan people from Xizang (Tibet Autonomous Region), 1.6% of a sample of Xibe people from Xinjiang, 3.2% of a sample of northern Hui from Ningxia, 9.4% of a sample of Hazak (Kazakhs) from Xinjiang, and rates of 24.0%, 22.2%, 35.2%, 29.2% in 4 different samples of Uyghurs from Xinjiang, 9.1% in a sample of Mongols from Inner Mongolia, 10% of a sample of Northern Han Chinese from Gansu and 8.9% of a sample of Northern Han from western Henan. A different subclade of R1 was also found in 1.5% of a sample of northern Hui from Ningxia.[99] in the same study there were no cases of R1a detected at all in 6 samples of Han Chinese in Yunnan, 1 sample of Han in Guangxi, 5 samples of Han in Guizhou, 2 samples of Han in Guangdong, 2 samples of Han in Fujian, 2 samples of Han in Zhejiang, 1 sample of Han in Shanghai, 1 samples of Han in Jiangxi, 2 samples of Han in Hunan, 1 sample of Han in Hubei, 2 samples of Han in Sichuan, 1 sample of Han in Chongqing, 3 samples of Han in Shandong, 5 samples of Han in Gansu, 3 samples of Han in Jilin and 2 samples of Han in Heilongjiang.[100] In a 2014 paper, R1a1a has been detected in 1.8% (2/110) of Chinese samples. These two samples (R-M17, R-M198, R-M434, R-M458 for both) belonged to Han individuals from Fujian and Shanxi provinces.[101] 40% of Salars, 45.2% of Tajiks of Xinjiang, 54.3% of Dongxiang, 60.6% of Tatars and 68.9% of Kyrgyz in Xinjiang in northwestern China tested in one sample had R1a1-M17. Bao'an (Bonan) had the most haplogroup diversity of 0.8946±0.0305 while the other ethnic minorities in northwestern China had a high haplogroup diversity like Central Asians, of 0.7602±0.0546.[102]

In Eastern Siberia, R1a1a is found among certain indigenous ethnic groups including Kamchatkans and Chukotkans, and peaking in Itel'man at 22%.[103]

West Asia

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 and Iran. The Shimar (Shammar) Bedouin tribe in Kuwait show the highest frequency in the Middle East at 43%.[104][105][106]

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 et al. 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, 19% among Azerbaijanis in West Azerbaijan, 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 Bandari people in Hormozgan and 25% among the Baloch people in Sistan and Baluchestan Province.[107]

Di Cristofaro et al. (2013) found haplogroup R1a in 9.68% (18/186) of a set of samples from Iran, though with a large variance ranging from 0% (0/18) in a sample of Iranians from Tehran to 25% (5/20) in a sample of Iranians from Khorasan and 27% (3/11) in a sample of Iranians of unknown provenance. All Iranian R1a individuals carried the M198 and M17 mutations except one individual in a sample of Iranians from Gilan (n=27), who was reported to belong to R1a-SRY1532.2(xM198, M17).[90]

Malyarchuk et al. (2013) found R1a1-SRY10831.2 in 20.8% (16/77) of a sample of Persians collected in the provinces of Khorasan and Kerman in eastern Iran, but they did not find any member of this haplogroup in a sample of 25 Kurds collected in the province of Kermanshah in western Iran.[91]

Further to the north of these Western Asian 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.[3]

Historic naming of R1a

Further information: Conversion table for Y chromosome haplogroups

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 et al. 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[citation needed] The discovery of M420 in 2009 has caused a reassignment of these phylogenetic names.(Underhill et al. 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.

Contrasting family trees for R1a, showing the evolution of understanding of this clade
2002 scheme proposed in (YCC 2002) 2009 scheme as per (Underhill et al. 2009) ISOGG tree as per January 2011[citation needed]
As M420 went undetected, M420 lineages were classified as either R1* or R1a (SRY1532.2, also known as SRY10831.2)
R1
 M173  
R1*

 All cases without M343 or SRY1532.2 (including a minority M420+ cases)

R1a
 SRY1532.2 
  (SRY10831.2)  

R1a* 

 
R1a1
 M17, M198 

 R1a1*

 M56 

 R1a1a

 M157 

 R1a1b

 M87, M204
M64.2

 
 R1a1c

R1b
M343

 sibling clade to R1a

After 2009, a new layer was inserted covering all old R1a, plus its closest known relatives
R1
 M173  
R1*

 All cases without M343 or M420 (smaller than old "R1a*")

R1a 
M420 

  R1a* All cases with M420 but without SRY1532.2

R1a1 
SRY1532.2 

  R1a1*(Old R1a*)

 R1a1a 
 M17, M198 

R1a1a*

M56
 

R1a1a1

M157
 

R1a1a2

 M64.2,..
 

R1a1a3

P98
 

R1a1a4

PK5
 

R1a1a5

M434
 

R1a1a6

 M458 
 

 R1a1a7*

 
M334 
 

 R1a1a7a

 Page68

R1a1a8

R1b
M343

 Sibling clade to R1a (same as before)

Latest information
R1
M173

R1* (As before)

R1a
M420

R1a* (As before)

R1a1
SRY1532.2

R1a1* (As before)

R1a1a
M17

R1a1a* (As before)

R1a1a1
M417, Page7

R1a1a1*

M56
 

R1a1a1a

M157
 

R1a1a1b

 M64.2,..
 

R1a1a1c

P98
 

R1a1a1d

PK5
 

R1a1a1e

M434
 

R1a1a1f

 Z283 
 

 R1a1a1g*

 M458 
 

 R1a1a1g1*

 
M334 
 

 R1a1a1g1a


L260 
 

 R1a1a1g1b

 Z280 
 

 R1a1a1g2*

 
P278.2 
 

 R1a1a1g2a


L365 
 

 R1a1a1g2b


L366 
 

 R1a1a1g2c


Z92 
 

 R1a1a1g2d

 Z284 
 

 R1a1a1g3*

 
P278.2 
 

 R1a1a1g3a

 Z93

 R1a1a1h*

 
L342.2 
 

 R1a1a1h1*

 
L657 
 

 R1a1a1h1a

R1b
M343

Sibling clade to R1a (same as before)

See also

Y-DNA R-M207 subclades

Y-DNA backbone tree

Notes

  1. ^ According to Family Tree,[who?] they diversified c. 5,000 years ago.[9]
  2. ^ See also: "'Heavily sex-biased' population dispersals into the Indian Subcontinent (Silva et al. 2017)". Eurogenes Blog. March 28, 2017.[self-published source?]
  3. ^ Kivisild et al. (2003): "Haplogroup R1a, previously associated with the putative Indo-Aryan invasion, was found at its highest frequency in Punjab but also at a relatively high frequency (26%) in the Chenchu tribe. This finding, together with the higher R1a-associated short tandem repeat diversity in India and Iran compared with Europe and central Asia, suggests that southern and western Asia might be the source of this haplogroup."[20]
  4. ^ Sahoo et al. (2006): "... one should expect to observe dramatically lower genetic variation among Indian Rla lineages. In fact, the opposite is true: the STR haplotype diversity on the background of R1a in Central Asia (and also in Eastern Europe) has already been shown to be lower than that in India (6). Rather, the high incidence of R1* and Rla throughout Central Asian European populations (without R2 and R* in most cases) is more parsimoniously explained by gene flow in the opposite direction, possibly with an early founder effect in South or West Asia.[24]
  5. ^ Sharma et al. (2009): "A peculiar observation of the highest frequency (up to 72.22%) of Y-haplogroup R1a1* in Brahmins hinted at its presence as a founder lineage for this caste group. Further, observation of R1a1* in different tribal population groups, existence of Y-haplogroup R1a* in ancestors and extended phylogenetic analyses of the pooled dataset of 530 Indians, 224 Pakistanis and 276 Central Asians and Eurasians bearing the R1a1* haplogroup supported the autochthonous origin of R1a1 lineage in India and a tribal link to Indian Brahmins. However, it is important to discover novel Y-chromosomal binary marker(s) for a higher resolution of R1a1* and confirm the present conclusions."[1]
  6. ^ for most of the history, the Indian subcontinent was the most populated region of the world
  7. ^ Sengupta et al. (2006): "The widespread geographic distribution of HG R1a1-M17 across Eurasia and the current absence of informative subdivisions defined by binary markers leave uncertain the geographic origin of HG R1a1-M17. However, the contour map of R1a1-M17 variance shows the highest variance in the northwestern region of India ... The question remains of how distinctive is the history of L1 relative to some or all of R1a1 and R2 representatives. This uncertainty neutralizes previous conclusions that the intrusion of HGs R1a1 and R2 from the northwest in Dravidian-speaking southern tribes is attributable to a single recent event. [R1a1 and R2] could have actually arrived in southern India from a southwestern Asian source region multiple times, with some episodes considerably earlier than others. Considerable archeological evidence exists regarding the presence of Mesolithic peoples in India (Kennedy 2000), some of whom could have entered the subcontinent from the northwest during the late Pleistocene epoch. The high variance of R1a1 in India (table 12), the spatial frequency distribution of R1a1 microsatellite variance clines (fig. 4), and expansion time (table 11) support this view."[21]
  8. ^ Semenov & Bulat (2016) refer to the following publications:
    1. Haak, Wolfgang (2015). "Massive migration from the steppe is a source for Indo-European languages in Europe". Nature. 522 (7555): 207–211. arXiv:1502.02783. Bibcode:2015Natur.522..207H. bioRxiv 10.1101/013433. doi:10.1038/NATURE14317. PMC 5048219. PMID 25731166.
    2. Mathieson, Iain (2015). "Eight thousand years of natural selection in Europe". bioRxiv 10.1101/016477.
    3. Chekunova Е.М., Yartseva N.V., Chekunov М.К., Мazurkevich А.N. The First Results of the Genotyping of the Aboriginals and Human Bone Remains of the Archeological Memorials of the Upper Podvin'e. // Archeology of the lake settlements of IV—II Thousands BC: The chronology of cultures and natural environment and climatic rhythms. Proceedings of the International Conference, Devoted to the 50-year Research of the Pile Settlements on the North-West of Russia. St. Petersburg, 13–15 November 2014.
    4. Jones, ER; Gonzalez-Fortes, G; Connell, S; Siska, V; Eriksson, A; Martiniano, R; McLaughlin, RL; Gallego Llorente, M; Cassidy, LM; Gamba, C; Meshveliani, T; Bar-Yosef, O; Müller, W; Belfer-Cohen, A; Matskevich, Z; Jakeli, N; Higham, TF; Currat, M; Lordkipanidze, D; Hofreiter, M; Manica, A; Pinhasi, R; Bradley, DG (2015). "Upper Palaeolithic genomes reveal deep roots of modern Eurasians". Nat Commun. 6: 8912. Bibcode:2015NatCo...6.8912J. doi:10.1038/ncomms9912. PMC 4660371. PMID 26567969.
  9. ^ Yet, Haak et al. also explicitly state: "a type of Near Eastern ancestry different from that which was introduced by early farmers".[clarification needed][33]
  10. ^ According to Family Tree DNA, L664 formed 4,700 ybp, that is, 2,700 BCE.[9]
  11. ^ Lazaridis, Twitter, 18 June 2016: "I1635 (Armenia_EBA) is R1b1-M415(xM269). We'll be sure to include in the revision. Thanks to the person who noticed! #ILovePreprints."[unreliable source?]
    See also "Big deal of 2016: the territory of present-day Iran cannot be the Indo-European homeland". Eurogenes Blog. November 26, 2016,[unreliable source?] for a discussion of the same topic.
  12. ^ See map for M780 distribution at Dieneke's Anthropology Blog, Major new article on the deep origins of Y-haplogroup R1a (Underhill et al. 2014)[37]
  13. ^ According to Family Tree DNA, M780 formed 4700 ybp.[9] This dating coincides with the eastward movement between 2800 and 2600 BCE of the Yamnaya culture into the region of the Poltavka culture, a predecessor of the Sintashta culture, from which the Indo-Iranians originated. M780 is concentrated in the Ganges Valley, the locus of the classic Vedic society.
  14. ^ Poznik et al. (2016) calculate with a generation time of 30 years; a generation time of 20 years yields other results.
  15. ^ "The evidence that the Steppe_MLBA [Middle to Late Bronze Age] cluster is a plausible source for the Steppe ancestry in South Asia is also supported by Y chromosome evidence, as haplogroup R1a which is of the Z93 subtype common in South Asia today [Underhill et al. (2014), Silva et al. (2017)] was of high frequency in Steppe_MLBA (68%) (16), but rare in Steppe_EMBA [Early to Middle Bronze Age] (absent in our data)."[38]
  16. ^ Балановский (2015), p. 208 (in Russian) Прежде всего, это преобладание в славянских популяциях дославянского субстрата — двух ассимилированных ими генетических компонентов – восточноевропейского для западных и восточных славян и южноевропейского для южных славян...Можно с осторожностью предположить, что ассимилированный субстратмог быть представлен по преимуществу балтоязычными популяциями. Действительно, археологические данные указыва ют на очень широкое распространение балтских групп перед началом расселения славян. Балтскийсубстрату славян (правда, наряду с финно-угорским) выявляли и антропологи. Полученные нами генетические данные — и на графиках генетических взаимоотношений, и по доле общих фрагментов генома — указывают, что современные балтские народы являются ближайшими генетически ми соседями восточных славян. При этом балты являются и лингвистически ближайшими род ственниками славян. И можно полагать, что к моменту ассимиляции их генофонд не так сильно отличался от генофонда начавших свое широкое расселение славян. Поэтому если предположить,что расселяющиеся на восток славяне ассимилировали по преимуществу балтов, это может объяснить и сходство современных славянских и балтских народов друг с другом, и их отличия от окружающих их не балто-славянских групп Европы...В работе высказывается осторожное предположение, что ассимилированный субстрат мог быть представлен по преимуществу балтоязычными популяциями. Действительно, археологические данные указывают на очень широкое распространение балтских групп перед началом расселения славян. Балтский субстрат у славян (правда, наряду с финно-угорским) выявляли и антропологи. Полученные в этой работе генетические данные — и на графиках генетических взаимоотношений, и по доле общих фрагментов генома — указывают, что современные балтские народы являются ближайшими генетическими соседями восточных славян.

References

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  4. ^ a b Underhill et al. 2014, p. 130.
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  13. ^ a b Wells 2001.
  14. ^ a b c Pamjav et al. 2012.
  15. ^ a b Haak et al. 2015.
  16. ^ a b Allentoft et al. 2015.
  17. ^ a b Mathieson et al. 2015.
  18. ^ a b Silva et al. 2017.
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Further reading

  • Gimbutas (1970). Indo-European and Indo-Europeans. Philadelphia, PA: Univ. of Pennsylvania Press. pp. 155–195.
  • Patrizia Malaspina; Andrey I. Kozlov; Fulvio Cruciani; Piero Santolamazza; Nejat Akar; Dimiter Kovatchev; Marina G. Kerimova; Juri Parik; Richard Villems; Rosana Scozzari; Andrea Novelletto (2003). "Analysis of Y-chromosome variation in modern populations at the European-Asian border" (PDF). In K. Boyle; C. Renfrew; M. Levine (eds.). Ancient interactions: east and west in Eurasia. McDonald Institute Monographs. Cambridge: Cambridge University Press. pp. 309–313.
  • Parpola, Asko (2015). The Roots of Hinduism. The early Aryans and the Indus Civilisation. Oxford University Press.
  • Sharma, S. (2007). "1344/T: The Autochthonous Origin and a Tribal Link of Indian Brahmins: Evaluation Through Molecular Genetic Markers". (PDF) (Abstract). p. 273. Archived from the original (PDF) on 2008-06-26.
  • Trautmann, Thomas (2005). The Aryan Debate. Oxford University Press.
  • Varzari, Alexander (2006). Population History of the Dniester-Carpathians: Evidence from Alu Insertion and Y-Chromosome Polymorphisms (PDF) (Dissertation). München: Ludwig-Maximilians-Universität.
  • Wells, Spencer (2002). The Journey of Man: A Genetic Odyssey. Princeton University Press. ISBN 978-0-691-11532-0.
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  • Karafet, T. M.; Mendez, F. L.; Meilerman, M. B.; Underhill, P. A.; Zegura, S. L.; Hammer, M. F. (2008). "New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree". Genome Research. 18 (5): 830–838. doi:10.1101/gr.7172008. PMC 2336805. PMID 18385274. See also Supplementary Material.
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External links

 
Wikimedia Commons has media related to Haplogroup R1a of Y-DNA.
DNA Tree
  • FTDNA R1a Y-chromosome Haplogroup Project
  • R1a1a1 and Subclades Y-DNA Project – Background Family Tree DNA R1a1a1
TMRCA
  • TMRCA = Time to Most Recent Common Ancestor
Various
  • Danish Demes Regional DNA Project: Y-DNA Haplogroup R1a 2008-08-20 at the Wayback Machine
  • Eurogenes Blog, The Poltovka outlier
  • Avotaynu Online, The Y-DNA Fingerprint of the Shpoler Zeida, a Tzaddik Who Touched the World

January 17, 2023
haplogroup, haplogroup, m420, human, chromosome, haplogroup, which, distributed, large, region, eurasia, extending, from, scandinavia, central, europe, southern, siberia, south, asia, possible, time, origin22, years, agopossible, place, origineastern, europe, . Haplogroup R1a or haplogroup R M420 is a human Y chromosome DNA haplogroup which is distributed in a large region in Eurasia extending from Scandinavia and Central Europe to southern Siberia and South Asia 3 2 Haplogroup R1aPossible time of origin22 000 YBP 1 to 25 000 2 years agoPossible place of originEastern Europe South Asia Central Asia or West AsiaAncestorHaplogroup R1DescendantsR1a Z282 R1a Z93Defining mutationsR1a L62 L63 L120 M420 M449 M511 M513R1a1a M17 M198 M512 M514 M515 L168 L449 L457 L566Highest frequenciesSee List of R1a frequency by populationMap showing frequency of R1a haplogroup in Europe While R1a originated c 22 000 1 to 25 000 2 years ago its subclade M417 R1a1a1 diversified c 5 800 years ago 4 The place of origin of the subclade plays a role in the debate about the origins of Proto Indo Europeans The SNP mutation R M420 was discovered after R M17 R1a1a 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 Contents 1 Origins 1 1 R1a origins 1 2 Diversification of R1a1a1 M417 and ancient migrations 1 2 1 Proposed steppe dispersal of R1a1a 1 2 2 Proposed South Asian origins 1 2 3 Proposed Yamnaya origins 1 2 4 Proposed Transcaucasia and West Asian origins and possible influence on Indus Valley Civilization 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 R M417 R1a1a1 2 7 1 R Z282 R1a1a1b1a Eastern Europe 2 7 1 1 R M458 R1a1a1b1a1 2 7 1 1 1 R L260 R1a1a1b1a1a 2 7 1 1 2 R M334 2 7 1 2 R1a1a1b1a2 S466 Z280 S204 Z91 2 7 1 2 1 R1a1a1b1a2b3 Gwozdz s Cluster K 2 7 1 2 2 R1a1a1b1a2b3a R L365 2 7 2 R1a1a1b2 R Z93 Asia 3 Geographic distribution of R1a1a 3 1 Pre Historical 3 2 Historical to recent Europe 3 3 Asia 3 3 1 Central Asia 3 3 2 South Asia 3 3 3 East Asia 3 3 4 West Asia 4 Historic naming of R1a 5 See also 5 1 Y DNA R M207 subclades 5 2 Y DNA backbone tree 6 Notes 7 References 8 Sources 9 Further reading 10 External linksOrigins EditR1a origins Edit The split of R1a M420 is computed to c 22 000 1 or 25 000 2 years ago which is the time of the last glacial maximum A 2014 study by Peter A Underhill et al using 16 244 individuals from over 126 populations from across Eurasia concluded that there was a compelling case for the Middle East possibly near present day Iran as the geographic origin of hg R1a 2 The ancient DNA record has shown the first R1a during the Mesolithic in Eastern Hunter Gatherers from Eastern Europe 5 6 and the earliest case of R among Upper Paleolithic Ancient North Eurasians 7 from which the Eastern Hunter Gatherers predominantly derive their ancestry 8 Diversification of R1a1a1 M417 and ancient migrations Edit R1a origins Underhill 2009 3 R1a1a origins Pamjav et al 2012 possible migration R1a to Baltic coast and R1a1a oldest expansion and highest frequency Underhill et al 2014 According to Underhill et al 2014 the downstream R1a M417 subclade diversified into Z282 and Z93 circa 5 800 years ago in the vicinity of Iran and Eastern Turkey 4 note 1 Even though R1a occurs as a Y chromosome haplogroup among various languages such as Slavic and Indo Iranian 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 civilization R1a shows a strong correlation with Indo European languages of Southern and Western Asia Central and Eastern Europe and to some extent Scandinavia 10 3 being most prevalent in Eastern Europe West Asia and South Asia In Europe Z282 is prevalent particularly while in Asia Z93 dominates The connection between Y DNA R M17 and the spread of Indo European languages was first noted by T Zerjal and colleagues in 1999 11 Proposed steppe dispersal of R1a1a Edit See also Indo European migrations Proto Indo Europeans and Indo Aryan migration Haplogroup R1a and related haplogroups Semino et al 2000 proposed Ukrainian origins and a postglacial spread of the R1a1 haplogroup during the Late Glacial Maximum subsequently magnified by the expansion of the Kurgan culture into Europe and eastward 12 Spencer Wells proposes Central Asian origins suggesting 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 13 According to Pamjav et al 2012 R1a1a diversified in the Eurasian Steppes or the Middle East and Caucasus region 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 14 Three genetic studies in 2015 gave support to the Kurgan theory of Gimbutas regarding the Indo European Urheimat According to those studies haplogroups R1b and R1a now the most common in Europe R1a is also common in South Asia would have expanded from the Pontic Caspian steppes along with the Indo European languages they also detected an autosomal component present in modern Europeans which was not present in Neolithic Europeans which would have been introduced with paternal lineages R1b and R1a as well as Indo European languages 15 16 17 Silva et al 2017 noted that R1a in South Asia most likely spread from a single Central Asian source pool there do seem to be at least three and probably more R1a founder clades within the Subcontinent consistent with multiple waves of arrival 18 According to Martin P Richards co author of Silva et al 2017 the prevalence of R1a in India was very powerful evidence for a substantial Bronze Age migration from central Asia that most likely brought Indo European speakers to India 19 note 2 Proposed South Asian origins Edit Kivisild et al 2003 have proposed either South or West Asia 20 note 3 while Mirabal et al 2009 see support for both South and Central Asia 10 Sharma et al 2009 showcased the existence of R1a in India beyond 18 000 years to possibly 44 000 years in origin 1 South Asian populations have the highest STR diversity within R1a1a 21 22 10 3 1 23 and subsequent older TMRCA datings citation needed and R1a1a is present among both higher Brahmin castes and lower castes although the frequency is higher among Brahmin castes Nevertheless the oldest TMRCA datings of the R1a haplogroup occur in the Saharia tribe a scheduled caste of the Bundelkhand region 1 23 From these findings some researchers have concluded that R1a1a originated in South Asia 22 1 note 4 note 5 excluding a more recent yet minor genetic influx from Indo European migrants in northwestern regions such as Afghanistan Balochistan Punjab and Kashmir 22 21 3 However this diversity and the subsequent older TMRCA datings can also be explained by the historically high population numbers note 6 which increases the likelihood of diversification and microsatellite variation 19 18 According to Sengupta et al 2006 R1a1 and R2 could have actually arrived in southern India from a southwestern Asian source region multiple times 21 note 7 However Sengupta also described in this article We found that the influence of Central Asia on the pre existing gene pool was minor The ages of accumulated microsatellite variation in the majority of Indian haplogroups exceed 10 000 15 000 years which attests to the antiquity of regional differentiation Therefore our data do not support models that invoke a pronounced recent genetic input from Central Asia to explain the observed genetic variation in South Asia This suggests that the origins of paternal haplogroup R1a point to the Indian subcontinent and not Central Asia Proposed Yamnaya origins Edit European middle Neolithic period Comb Ware culture c 4200 c 2000 BCE Corded Ware culture c 2900 c 2350 BCE David Anthony considers the Yamnaya culture to be the Indo European Urheimat 25 26 According to Haak et al 2015 a massive migration from the Yamnaya culture northwards took place c 2 500 BCE accounting for 75 of the genetic ancestry of the Corded Ware culture noting that R1a and R1b may have spread into Europe from the East after 3 000 BCE 27 Yet all their seven Yamnaya samples belonged to the R1b M269 subclade 27 but no R1a1a has been found in their Yamnaya samples This raises the question where the R1a1a in the Corded Ware culture came from if it was not from the Yamnaya culture 28 According to Marc Haber the absence of haplogroup R1a M458 in Afghanistan does not support a Pontic Caspian steppe origin for the R1a lineages in modern Central Asian populations 29 According to Leo Klejn the absence of haplogroup R1a in Yamnaya remains despite its presence in Eneolithic Samara and Eastern Hunter Gatherer populations makes it unlikely that Europeans inherited haplogroup R1a from Yamnaya 30 Archaeologist Barry Cunliffe has said that the absence of haplogroup R1a in Yamnaya specimens is a major weakness in Haak s proposal that R1a has a Yamnaya origin 31 Semenov amp Bulat 2016 do argue for a Yamnaya origin of R1a1a in the Corded Ware culture noting that several publications point to the presence of R1a1 in the Comb Ware culture 32 note 8 Proposed Transcaucasia and West Asian origins and possible influence on Indus Valley Civilization Edit See also Kura Araxes culture Uruk period and Indus Valley Civilisation Possible near Eastern Dravidian origins Haak et al 2015 found that part of the Yamnaya ancestry derived from the Middle East and that neolithic techniques probably arrived at the Yamnaya culture from the Balkans note 9 The Rossen culture 4 600 4 300 BC which was situated on Germany and predates the Corded Ware culture an old subclade of R1a namely L664 can still be found note 10 Part of the South Asian genetic ancestry derives from west Eurasian populations and some researchers have implied that Z93 may have come to India via Iran 34 and expanded there during the Indus Valley civilization 2 35 Mascarenhas et al 2015 proposed that the roots of Z93 lie in West Asia and proposed that Z93 and L342 2 expanded in a southeasterly direction from Transcaucasia into South Asia 34 noting that such an expansion 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 34 Yet Lazaridis noted that sample I1635 of Lazaridis et al 2016 their Armenian Kura Araxes sample carried Y haplogroup R1b1 M415 xM269 note 11 also called R1b1a1b CTS3187 36 unreliable source According to Underhill et al 2014 the diversification of Z93 and the early urbanization within the Indus Valley occurred at 5 600 years ago and the geographic distribution of R1a M780 Figure 3d note 12 may reflect this 2 note 13 Poznik et al 2016 note that striking expansions occurred within R1a Z93 at c 4 500 4 000 years ago which predates by a few centuries the collapse of the Indus Valley Civilisation 35 note 14 However according to Narasimhan et al 2018 steppe pastoralists are a likely source for R1a in India 38 note 15 Phylogeny EditThe 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 such as R1a1 in relatively recent but not the latest literature Topology Edit The topology of R1a is as follows codes in brackets non isogg codes 9 39 verification needed 40 2 41 Tatiana et al 2014 rapid diversification process of K M526 likely occurred in Southeast Asia with subsequent westward expansions of the ancestors of haplogroups R and Q 42 P P295 PF5866 S8 also known as K2b2 R R M207 40 9 R R1 R M173 R1 40 R1a M420 40 Eastern Europe Asia 2 R1a 9 R1a1 40 M459 PF6235 40 SRY1532 2 SRY10831 2 40 R1a1 M459 40 9 R1a1a M17 M198 40 R1a1a1 M417 page7 40 R1a1a1a CTS7083 L664 S298 40 R1a1a1b S224 Z645 S441 Z647 40 R1a1a1b1 PF6217 S339 Z283 40 R1a1a1b1a Z282 40 R1a1a1a Z282 43 R1a1a1b1a1 40 The old topological code is R1a1a1b which is outdated and might lead to some confusion 43 M458 40 43 R1a1a1g M458 41 R1a1a1g 41 R1a1a1g1 M334 41 R1a1a1b1a1a L260 S222 40 R1a1a1g2 41 R1a1a1b1a2 40 S466 Z280 S204 Z91 40 R1a1a1b1a2a 40 R1a1a1b1a2b CTS1211 40 R1a1a1c M558 43 R CTS1211 V2803 CTS3607 S3363 M558 CTS1211 S3357 Y34 FGC36457 9 R1a1a1b1a2b3 M417 Z645 Z283 Z282 Z280 CTS1211 CTS3402 Y33 CTS3318 Y2613 Gwozdz s Cluster K 39 verification needed R1a1a1b1a2b3a L365 S468 40 R1a1a1b1a3 Z284 40 R1a1a1a1 Z284 43 R1a1a1b2 F992 S202 Z93 40 R1a1a2 Z93 M746 43 R1a1a1b2a F3105 S340 Z94 L342 2 S278 2 40 R1a1b2a Z95 43 R Z94 Z94 F3105 S340 Z95 F3568 9 R Z2124 Z2121 S3410 Z2124 9 R1a1b2a Z2125 43 R1a1b2a M434 43 R1a1a1f M434 41 R1a1b2a M204 43 R1a1b2a1 M560 43 R1a1b2a2 M780 L657 43 India 2 R1a1b2a3 Z2122 M582 43 R1a1a1c M64 2 M87 M204 41 R1a1a1d P98 41 R1a1a1d2a 44 R1a1a1e PK5 41 R1b M343 Western Europe R2 India Haplogroup R Edit Haplogroup R phylogeny R M207 R1 M173 M420 R1a M343 R1b M173 xM420 M343 R1 R2 M479 R M207 xM173 M479 R M173 R1 Edit R1a is distinguished by several unique markers including the M420 mutation It 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 R1a Edit 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 3 45 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 3 This paragroup is now known as R1a2 R YP4141 It then has two branches R1a2a R YP5018 and R1a2b R YP4132 R SRY1532 2 R1a1 Edit R1a1 is defined by SRY1532 2 or SRY10831 2 understood to always include SRY10831 2 M448 L122 M459 and M516 3 46 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 al 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 3 Sahoo et al 2006 reported R SRY1532 2 for 1 15 Himachal Pradesh Rajput samples 22 R M17 M198 R1a1a Edit The following SNPs are associated with R1a1a SNP Mutation Y position NCBI36 Y position GRCh37 RefSNP IDM17 INS G 20192556 21733168 rs3908M198 C gt T 13540146 15030752 rs2020857M512 C gt T 14824547 16315153 rs17222146M514 C gt T 17884688 19375294 rs17315926M515 T gt A 12564623 14054623 rs17221601L168 A gt G 14711571 16202177 L449 C gt T 21376144 22966756 L457 G gt A 14946266 16436872 rs113195541L566 C gt T R M417 R1a1a1 Edit R1a1a1 R M417 is the most widely found subclade in two variations which are found respectively in Europe R1a1a1b1 R Z282 R1a1a1a R Z282 Underhill 2014 2 and Central and South Asia R1a1a1b2 R Z93 R1a1a2 R Z93 Underhill 2014 2 R Z282 R1a1a1b1a Eastern Europe Edit This large subclade appears to encompass most of the R1a1a found in Europe 14 R1a1a1b1a R1a1a1a Underhill 2014 R Z282 occurs in northern Ukraine Belarus and Russia at a frequency of c 20 2 R1a1a1b1a3 R1a1a1a1 Underhill 2014 R Z284 occurs in Northwest Europe and peaks at c 20 in Norway 2 R1a1a1c M64 2 M87 M204 is apparently rare it was found in 1 of 117 males typed in southern Iran 47 R M458 R1a1a1b1a1 Edit Frequency distribution of R M458 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 3 M458 was found in one skeleton from a 14th century grave field in Usedom Mecklenburg Vorpommern Germany 48 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 R L260 R1a1a1b1a1a Edit 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 et al 2002 In 2010 it was verified to be a haplogroup identified by its own mutation SNP 49 It apparently accounts for about 8 of Polish men making it the most common subclade in Poland Outside of Poland it is less common 50 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 51 R M334 Edit R M334 R1a1a1g1 41 a subclade of R1a1a1g M458 41 c q R1a1a1b1a1 M458 40 was found by Underhill et al 2009 only in one Estonian man and may define a very recently founded and small clade 3 R1a1a1b1a2 S466 Z280 S204 Z91 Edit R1a1a1b1a2b3 Gwozdz s Cluster K Edit R1a1a1b1a2b3 M417 Z645 Z283 Z282 Z280 CTS1211 CTS3402 Y33 CTS3318 Y2613 Gwozdz s Cluster K 39 verification needed is a STR based group that is R M17 xM458 This cluster is common in Poland but not exclusive to Poland 51 R1a1a1b1a2b3a R L365 Edit R1a1a1b1a2b3a R L365 40 was early called Cluster G citation needed R1a1a1b2 R Z93 Asia Edit Relative frequency of R M434 to R M17 Region People N R M17 R M434Number 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 5 Table only shows positive sets from N 3667 derived from 60 Eurasian populations sample 3 This large subclade appears to encompass most of the R1a1a found in Asia being related to Indo European migrations including Scythians Indo Aryan migrations and so on 14 R Z93 or R1a1a1b2 R1a1a2 in Underhill 2014 is most common gt 30 in the South Siberian Altai region of Russia cropping up in Kyrgyzstan 6 and in all Iranian populations 1 8 2 R Z2125 occurs at highest frequencies in Kyrgyzstan and in Afghan Pashtuns gt 40 At a frequency of gt 10 it is also observed in other Afghan ethnic groups and in some populations in the Caucasus and Iran 2 R M434 is a subclade of Z2125 It was detected in 14 people out of 3667 people tested all in a restricted geographical range from Pakistan to Oman This likely reflects a recent mutation event in Pakistan 3 R M560 is very rare and was only observed in four samples two Burushaski speakers north Pakistan one Hazara Afghanistan and one Iranian Azerbaijani 2 R M780 occurs at high frequency in South Asia India Pakistan Afghanistan and the Himalayas The group also occurs at gt 3 in some Iranian populations and is present at gt 30 in Roma from Croatia and Hungary 2 Geographic distribution of R1a1a Edit Distribution of R1a purple and R1b red Pre Historical Edit In Mesolithic Europe R1a is characteristic of Eastern Hunter Gatherers EHGs 52 A male EHG of the Veretye culture buried at Peschanitsa near Lake Lacha in Arkhangelsk Oblast Russia c 10 700 BCE was found to be a carrier of the paternal haplogroup R1a5 YP1301 and the maternal haplogroup U4a 53 54 52 A Mesolithic male from Karelia c 8 800 BCE to 7950 BCE has been found to be carrying haplogroup R1a 55 A Mesolithic male buried at Deriivka c 7000 BCE to 6700 BCE carried the paternal haplogroup R1a and the maternal U5a2a 17 Another male from Karelia from c 5 500 to 5 000 BC who was considered an EHG carried haplogroup R1a 15 A male from the Comb Ceramic culture in Kudrukula c 5 900 BCE to 3 800 BCE has been determined to be a carrier of R1a and the maternal U2e1 56 According to archaeologist David Anthony the paternal R1a Z93 was found at Alexandria Ukraine c 4000 BCE Sredny Stog culture the earliest known sample to show the genetic adaptation to lactase persistence 13910 T 57 R1a has been found in the Corded Ware culture 58 59 in which it is predominant 60 Examined males of the Bronze Age Fatyanovo culture belong entirely to R1a specifically subclade R1a Z93 52 53 61 Haplogroup R1a has later been found in ancient fossils associated with the Urnfield culture 62 as well as the burial of the remains of the Sintashta 16 Andronovo 63 the Pazyryk 64 Tagar 63 Tashtyk 63 and Srubnaya cultures the inhabitants of ancient Tanais 65 in the Tarim mummies 66 and the aristocracy of Xiongnu 67 The skeletal 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 58 Historical to recent Europe Edit In Europe the R1a1 sub clade is found at highest levels among peoples of Central and Eastern European descent with results ranging from 35 to 65 among Czechs Hungarians Poles Slovaks western Ukrainians particularly Rusyns Belarusians Moldovans and Russians 68 69 12 In the Baltics R1a1a frequencies decrease from Lithuania 45 to Estonia around 30 70 71 72 12 73 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 74 75 Vikings and Normans may have also carried the R1a1a lineage further out accounting for at least part of the small presence in the British Isles the Canary Islands and Sicily 76 77 In East Germany Haplogroup R1a1a averages between 20 and 30 with a peak in Rostock at 31 3 it 78 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 79 better source needed The Balkans shows wide variation between areas with significant levels of R1a1a for example 36 39 in Slovenia 80 27 34 in Croatia 71 81 82 83 84 and over 30 in Greek Macedonia but less than 10 in Albania Kosovo and parts of Greece south of Olympus gorge 85 72 12 R1a is virtually composed only of the Z284 subclade in Scandinavia In Slovenia the main subclade is Z282 Z280 and M458 although the Z284 subclade was found in one sample of a Slovenian There is a negligible representation of Z93 in each region other than Turkey 2 West Slavs and Hungarians are characterized by a high frequency of the subclade M458 and a low Z92 a subclade of Z280 Hundreds of Slovenian samples and Czechs lack the Z92 subclade of Z280 while Poles Slovaks Croats and Hungarians only show a very low frequency of Z92 2 The Balts East Slavs Serbs Macedonians Bulgarians and Romanians demonstrate a ratio Z280 gt M458 and a high up to a prevailing share of Z92 2 Balts and East Slavs have the same subclades and similar frequencies in a more detailed phylogeny of the subclades 86 87 The Russian geneticist Oleg Balanovsky speculated that there is a predominance of the assimilated pre Slavic substrate in the genetics of East and West Slavic populations according to him the common genetic structure which contrasts East Slavs and Balts from other populations may suggest the explanation that the pre Slavic substrate of the East Slavs consisted most significantly of Baltic speakers which at one point predated the Slavs in the cultures of the Eurasian steppe according to archaeological and toponymic references note 16 Asia Edit Central Asia Edit Zerjal et al 2002 found R1a1a in 64 of a sample of the Tajiks of Tajikistan and 63 of a sample of the Kyrgyz of Kyrgyzstan 88 Haber et al 2012 found R1a1a M17 in 26 0 53 204 of a set of samples from Afghanistan including 60 3 5 of a sample of Nuristanis 51 0 25 49 of a sample of Pashtuns 30 4 17 56 of a sample of Tajiks 17 6 3 17 of a sample of Uzbeks 6 7 4 60 of a sample of Hazaras and in the only sampled Turkmen individual 89 Di Cristofaro et al 2013 found R1a1a M198 M17 in 56 3 49 87 of a pair of samples of Pashtuns from Afghanistan including 20 34 or 58 8 of a sample of Pashtuns from Baghlan and 29 53 or 54 7 of a sample of Pashtuns from Kunduz 29 1 37 127 of a pool of samples of Uzbeks from Afghanistan including 28 94 or 29 8 of a sample of Uzbeks from Jawzjan 8 28 or 28 6 of a sample of Uzbeks from Sar e Pol and 1 5 or 20 of a sample of Uzbeks from Balkh 27 5 39 142 of a pool of samples of Tajiks from Afghanistan including 22 54 or 40 7 of a sample of Tajiks from Balkh 9 35 or 25 7 of a sample of Tajiks from Takhar 4 16 or 25 0 of a sample of Tajiks from Samangan and 4 37 or 10 8 of a sample of Tajiks from Badakhshan 16 2 12 74 of a sample of Turkmens from Jawzjan and 9 1 7 77 of a pair of samples of Hazara from Afghanistan including 7 69 or 10 1 of a sample of Hazara from Bamiyan and 0 8 or 0 of a sample of Hazara from Balkh 90 Malyarchuk et al 2013 found R1a1 SRY10831 2 in 30 0 12 40 of a sample of Tajiks from Tajikistan 91 Ashirbekov et al 2017 found R1a M198 in 6 03 78 1294 of a set of samples of Kazakhs from Kazakhstan R1a M198 was observed with greater than average frequency in the study s samples of the following Kazakh tribes 13 41 31 7 of a sample of Suan 8 29 27 6 of a sample of Oshaqty 6 30 20 0 of a sample of Qozha 4 29 13 8 of a sample of Qypshaq 1 8 12 5 of a sample of Tore 9 86 10 5 of a sample of Jetyru 4 50 8 0 of a sample of Argyn 1 13 7 7 of a sample of Shanyshqyly 8 122 6 6 of a sample of Alimuly 3 46 6 5 of a sample of Alban R1a M198 also was observed in 5 42 11 9 of a sample of Kazakhs of unreported tribal affiliation 92 South Asia Edit In South Asia R1a1a has often been observed in a number of demographic groups 22 21 In India high frequencies of this haplogroup is observed in West Bengal Brahmins 72 21 to the east Gujarat Lohanas 60 3 to the west Khatris 67 3 in the north and Iyengar Brahmins 31 21 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 Kota 22 58 93 and the Kallar of Tamil Nadu suggesting that R1a1a is widespread in Tribal Southern Indians 20 Besides these studies show high percentages in regionally diverse groups such as Manipuris 50 3 to the extreme North East and among Punjabis 47 20 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 3 Among the Sinhalese of Sri Lanka 23 were found to be R1a1a R SRY1532 positive 94 Hindus of Chitwan District in the Terai region Nepal show it at 69 95 East Asia Edit The frequency of R1a1a is comparatively low among some Turkic speaking groups like 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 13 96 97 A Chinese paper published in 2018 found R1a Z94 in 38 5 15 39 of a sample of Keriyalik Uyghurs from Darya Boyi Darya Boye Village Yutian County Xinjiang 于田县达里雅布依乡 R1a Z93 in 28 9 22 76 of a sample of Dolan Uyghurs from Horiqol township Awat County Xinjiang 阿瓦提县乌鲁却勒镇 and R1a Z93 in 6 3 4 64 of a sample of Loplik Uyghurs from Karquga Qarchugha Village Yuli County Xinjiang 尉犁县喀尔曲尕乡 R1a xZ93 was observed only in one of 76 Dolan Uyghurs 98 Note that Darya Boyi Village is located in a remote oasis formed by the Keriya River in the Taklamakan Desert A 2011 Y DNA study found Y dna R1a1 in 10 of a sample of southern Hui people from Yunnan 1 6 of a sample of Tibetan people from Xizang Tibet Autonomous Region 1 6 of a sample of Xibe people from Xinjiang 3 2 of a sample of northern Hui from Ningxia 9 4 of a sample of Hazak Kazakhs from Xinjiang and rates of 24 0 22 2 35 2 29 2 in 4 different samples of Uyghurs from Xinjiang 9 1 in a sample of Mongols from Inner Mongolia 10 of a sample of Northern Han Chinese from Gansu and 8 9 of a sample of Northern Han from western Henan A different subclade of R1 was also found in 1 5 of a sample of northern Hui from Ningxia 99 in the same study there were no cases of R1a detected at all in 6 samples of Han Chinese in Yunnan 1 sample of Han in Guangxi 5 samples of Han in Guizhou 2 samples of Han in Guangdong 2 samples of Han in Fujian 2 samples of Han in Zhejiang 1 sample of Han in Shanghai 1 samples of Han in Jiangxi 2 samples of Han in Hunan 1 sample of Han in Hubei 2 samples of Han in Sichuan 1 sample of Han in Chongqing 3 samples of Han in Shandong 5 samples of Han in Gansu 3 samples of Han in Jilin and 2 samples of Han in Heilongjiang 100 In a 2014 paper R1a1a has been detected in 1 8 2 110 of Chinese samples These two samples R M17 R M198 R M434 R M458 for both belonged to Han individuals from Fujian and Shanxi provinces 101 40 of Salars 45 2 of Tajiks of Xinjiang 54 3 of Dongxiang 60 6 of Tatars and 68 9 of Kyrgyz in Xinjiang in northwestern China tested in one sample had R1a1 M17 Bao an Bonan had the most haplogroup diversity of 0 8946 0 0305 while the other ethnic minorities in northwestern China had a high haplogroup diversity like Central Asians of 0 7602 0 0546 102 In Eastern Siberia R1a1a is found among certain indigenous ethnic groups including Kamchatkans and Chukotkans and peaking in Itel man at 22 103 West Asia Edit 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 and Iran The Shimar Shammar Bedouin tribe in Kuwait show the highest frequency in the Middle East at 43 104 105 106 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 et al 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 19 among Azerbaijanis in West Azerbaijan 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 Bandari people in Hormozgan and 25 among the Baloch people in Sistan and Baluchestan Province 107 Di Cristofaro et al 2013 found haplogroup R1a in 9 68 18 186 of a set of samples from Iran though with a large variance ranging from 0 0 18 in a sample of Iranians from Tehran to 25 5 20 in a sample of Iranians from Khorasan and 27 3 11 in a sample of Iranians of unknown provenance All Iranian R1a individuals carried the M198 and M17 mutations except one individual in a sample of Iranians from Gilan n 27 who was reported to belong to R1a SRY1532 2 xM198 M17 90 Malyarchuk et al 2013 found R1a1 SRY10831 2 in 20 8 16 77 of a sample of Persians collected in the provinces of Khorasan and Kerman in eastern Iran but they did not find any member of this haplogroup in a sample of 25 Kurds collected in the province of Kermanshah in western Iran 91 Further to the north of these Western Asian 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 3 Historic naming of R1a EditFurther information Conversion table for Y chromosome haplogroups 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 et al 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 citation needed The discovery of M420 in 2009 has caused a reassignment of these phylogenetic names Underhill et al 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 Contrasting family trees for R1a showing the evolution of understanding of this clade 2002 scheme proposed in YCC 2002 2009 scheme as per Underhill et al 2009 ISOGG tree as per January 2011 citation needed As M420 went undetected M420 lineages were classified as either R1 or R1a SRY1532 2 also known as SRY10831 2 R1 M173 R1 All cases without M343 or SRY1532 2 including a minority M420 cases R1a SRY1532 2 SRY10831 2 R1a R1a1 M17 M198 R1a1 M56 R1a1a M157 R1a1b M87 M204M64 2 R1a1cR1bM343 sibling clade to R1a After 2009 a new layer was inserted covering all old R1a plus its closest known relativesR1 M173 R1 All cases without M343 or M420 smaller than old R1a R1a M420 R1a All cases with M420 but without SRY1532 2R1a1 SRY1532 2 R1a1 Old R1a R1a1a M17 M198 R1a1a M56 R1a1a1M157 R1a1a2 M64 2 R1a1a3P98 R1a1a4PK5 R1a1a5M434 R1a1a6 M458 R1a1a7 M334 R1a1a7a Page68 R1a1a8R1bM343 Sibling clade to R1a same as before Latest informationR1M173 R1 As before R1aM420 R1a As before R1a1SRY1532 2 R1a1 As before R1a1aM17 R1a1a As before R1a1a1M417 Page7 R1a1a1 M56 R1a1a1aM157 R1a1a1b M64 2 R1a1a1cP98 R1a1a1dPK5 R1a1a1eM434 R1a1a1f Z283 R1a1a1g M458 R1a1a1g1 M334 R1a1a1g1aL260 R1a1a1g1b Z280 R1a1a1g2 P278 2 R1a1a1g2aL365 R1a1a1g2bL366 R1a1a1g2cZ92 R1a1a1g2d Z284 R1a1a1g3 P278 2 R1a1a1g3a Z93 R1a1a1h L342 2 R1a1a1h1 L657 R1a1a1h1aR1bM343 Sibling clade to R1a same as before See also EditList of R1a frequency by population Genetic history of Europe Genetics and archaeogenetics of South Asia Haplogroup Q M242 Y DNA Human Y chromosome DNA haplogroups Neanderthal Y chromosome DNA haplogroups Y chromosome haplogroups in populations of the world Y DNA R M207 subclades Edit R L21 R L295 R M124 R M167 R M17 R M173 R M207 R M342 R M420 R M479 R U106 Y DNA backbone tree EditNotes Edit According to Family Tree who they diversified c 5 000 years ago 9 See also Heavily sex biased population dispersals into the Indian Subcontinent Silva et al 2017 Eurogenes Blog March 28 2017 self published source Kivisild et al 2003 Haplogroup R1a previously associated with the putative Indo Aryan invasion was found at its highest frequency in Punjab but also at a relatively high frequency 26 in the Chenchu tribe This finding together with the higher R1a associated short tandem repeat diversity in India and Iran compared with Europe and central Asia suggests that southern and western Asia might be the source of this haplogroup 20 Sahoo et al 2006 one should expect to observe dramatically lower genetic variation among Indian Rla lineages In fact the opposite is true the STR haplotype diversity on the background of R1a in Central Asia and also in Eastern Europe has already been shown to be lower than that in India 6 Rather the high incidence of R1 and Rla throughout Central Asian European populations without R2 and R in most cases is more parsimoniously explained by gene flow in the opposite direction possibly with an early founder effect in South or West Asia 24 Sharma et al 2009 A peculiar observation of the highest frequency up to 72 22 of Y haplogroup R1a1 in Brahmins hinted at its presence as a founder lineage for this caste group Further observation of R1a1 in different tribal population groups existence of Y haplogroup R1a in ancestors and extended phylogenetic analyses of the pooled dataset of 530 Indians 224 Pakistanis and 276 Central Asians and Eurasians bearing the R1a1 haplogroup supported the autochthonous origin of R1a1 lineage in India and a tribal link to Indian Brahmins However it is important to discover novel Y chromosomal binary marker s for a higher resolution of R1a1 and confirm the present conclusions 1 for most of the history the Indian subcontinent was the most populated region of the world Sengupta et al 2006 The widespread geographic distribution of HG R1a1 M17 across Eurasia and the current absence of informative subdivisions defined by binary markers leave uncertain the geographic origin of HG R1a1 M17 However the contour map of R1a1 M17 variance shows the highest variance in the northwestern region of India The question remains of how distinctive is the history of L1 relative to some or all of R1a1 and R2 representatives This uncertainty neutralizes previous conclusions that the intrusion of HGs R1a1 and R2 from the northwest in Dravidian speaking southern tribes is attributable to a single recent event R1a1 and R2 could have actually arrived in southern India from a southwestern Asian source region multiple times with some episodes considerably earlier than others Considerable archeological evidence exists regarding the presence of Mesolithic peoples in India Kennedy 2000 some of whom could have entered the subcontinent from the northwest during the late Pleistocene epoch The high variance of R1a1 in India table 12 the spatial frequency distribution of R1a1 microsatellite variance clines fig 4 and expansion time table 11 support this view 21 Semenov amp Bulat 2016 refer to the following publications Haak Wolfgang 2015 Massive migration from the steppe is a source for Indo European languages in Europe Nature 522 7555 207 211 arXiv 1502 02783 Bibcode 2015Natur 522 207H bioRxiv 10 1101 013433 doi 10 1038 NATURE14317 PMC 5048219 PMID 25731166 Mathieson Iain 2015 Eight thousand years of natural selection in Europe bioRxiv 10 1101 016477 Chekunova E M Yartseva N V Chekunov M K Mazurkevich A N The First Results of the Genotyping of the Aboriginals and Human Bone Remains of the Archeological Memorials of the Upper Podvin e Archeology of the lake settlements of IV II Thousands BC The chronology of cultures and natural environment and climatic rhythms Proceedings of the International Conference Devoted to the 50 year Research of the Pile Settlements on the North West of Russia St Petersburg 13 15 November 2014 Jones ER Gonzalez Fortes G Connell S Siska V Eriksson A Martiniano R McLaughlin RL Gallego Llorente M Cassidy LM Gamba C Meshveliani T Bar Yosef O Muller W Belfer Cohen A Matskevich Z Jakeli N Higham TF Currat M Lordkipanidze D Hofreiter M Manica A Pinhasi R Bradley DG 2015 Upper Palaeolithic genomes reveal deep roots of modern Eurasians Nat Commun 6 8912 Bibcode 2015NatCo 6 8912J doi 10 1038 ncomms9912 PMC 4660371 PMID 26567969 Yet Haak et al also explicitly state a type of Near Eastern ancestry different from that which was introduced by early farmers clarification needed 33 According to Family Tree DNA L664 formed 4 700 ybp that is 2 700 BCE 9 Lazaridis Twitter 18 June 2016 I1635 Armenia EBA is R1b1 M415 xM269 We ll be sure to include in the revision Thanks to the person who noticed ILovePreprints unreliable source See also Big deal of 2016 the territory of present day Iran cannot be the Indo European homeland Eurogenes Blog November 26 2016 unreliable source for a discussion of the same topic See map for M780 distribution at Dieneke s Anthropology Blog Major new article on the deep origins of Y haplogroup R1a Underhill et al 2014 37 According to Family Tree DNA M780 formed 4700 ybp 9 This dating coincides with the eastward movement between 2800 and 2600 BCE of the Yamnaya culture into the region of the Poltavka culture a predecessor of the Sintashta culture from which the Indo Iranians originated M780 is concentrated in the Ganges Valley the locus of the classic Vedic society Poznik et al 2016 calculate with a generation time of 30 years a generation time of 20 years yields other results The evidence that the Steppe MLBA Middle to Late Bronze Age cluster is a plausible source for the Steppe ancestry in South Asia is also supported by Y chromosome evidence as haplogroup R1a which is of the Z93 subtype common in South Asia today Underhill et al 2014 Silva et al 2017 was of high frequency in Steppe MLBA 68 16 but rare in Steppe EMBA Early to Middle Bronze Age absent in our data 38 Balanovskij 2015 p 208 in Russian Prezhde vsego eto preobladanie v slavyanskih populyaciyah doslavyanskogo substrata dvuh assimilirovannyh imi geneticheskih komponentov vostochnoevropejskogo dlya zapadnyh i vostochnyh slavyan i yuzhnoevropejskogo dlya yuzhnyh slavyan Mozhno s ostorozhnostyu predpolozhit chto assimilirovannyj substratmog byt predstavlen po preimushestvu baltoyazychnymi populyaciyami Dejstvitelno arheologicheskie dannye ukazyva yut na ochen shirokoe rasprostranenie baltskih grupp pered nachalom rasseleniya slavyan Baltskijsubstratu slavyan pravda naryadu s finno ugorskim vyyavlyali i antropologi Poluchennye nami geneticheskie dannye i na grafikah geneticheskih vzaimootnoshenij i po dole obshih fragmentov genoma ukazyvayut chto sovremennye baltskie narody yavlyayutsya blizhajshimi geneticheski mi sosedyami vostochnyh slavyan Pri etom balty yavlyayutsya i lingvisticheski blizhajshimi rod stvennikami slavyan I mozhno polagat chto k momentu assimilyacii ih genofond ne tak silno otlichalsya ot genofonda nachavshih svoe shirokoe rasselenie slavyan Poetomu esli predpolozhit chto rasselyayushiesya na vostok slavyane assimilirovali po preimushestvu baltov eto mozhet obyasnit i shodstvo sovremennyh slavyanskih i baltskih narodov drug s drugom i ih otlichiya ot okruzhayushih ih ne balto slavyanskih grupp Evropy V rabote vyskazyvaetsya ostorozhnoe predpolozhenie chto assimilirovannyj substrat mog byt predstavlen po preimushestvu baltoyazychnymi populyaciyami Dejstvitelno arheologicheskie dannye ukazyvayut na ochen shirokoe rasprostranenie baltskih grupp pered nachalom rasseleniya slavyan Baltskij substrat u slavyan pravda naryadu s finno ugorskim vyyavlyali i antropologi Poluchennye v etoj rabote geneticheskie dannye i na grafikah geneticheskih vzaimootnoshenij i po dole obshih fragmentov genoma ukazyvayut chto sovremennye baltskie narody yavlyayutsya blizhajshimi geneticheskimi sosedyami vostochnyh slavyan References Edit a b c d e f g h Sharma et al 2009 a b c d e f g h i j k l m n o p q r s t u Underhill et al 2014 a b c d e f g h i j k l m n o p q r Underhill et al 2009 a b Underhill et al 2014 p 130 Saag Lehti Vasilyev Sergey V Varul Liivi Kosorukova Natalia V Gerasimov Dmitri V Oshibkina Svetlana V Griffith Samuel J Solnik Anu Saag Lauri D Atanasio Eugenia Metspalu Ene January 2021 Genetic ancestry changes in Stone to Bronze Age transition in the East European plain Science 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tandem repeats with application to human population divergence time American Journal of Human Genetics 74 1 50 61 doi 10 1086 380911 PMC 1181912 PMID 14691732 G Mrsic et al 2012 Croatian national reference Y STR haplotype database Molecular Biology Reports 39 7 7727 41 doi 10 1007 s11033 012 1610 3 PMID 22391654 S2CID 18011987 J Sarac et al 2016 Genetic heritage of Croatians in the Southeastern European gene pool Y chromosome analysis of the Croatian continental and Island population American Journal of Human Biology 28 6 837 845 doi 10 1002 ajhb 22876 PMID 27279290 S2CID 25873634 External links Edit Wikimedia Commons has media related to Haplogroup R1a of Y DNA DNA TreeFTDNA R1a Y chromosome Haplogroup Project R1a1a1 and Subclades Y DNA Project Background Family Tree DNA R1a1a1TMRCATMRCA Time to Most Recent Common AncestorVariousDanish Demes Regional DNA Project Y DNA Haplogroup R1a Archived 2008 08 20 at the Wayback Machine Eurogenes Blog The Poltovka outlier Avotaynu Online The Y DNA Fingerprint of the Shpoler Zeida a Tzaddik Who Touched the World Retrieved from https en wikipedia org w index php title Haplogroup R1a amp oldid 1131548081, wikipedia, wiki, book, books, library,

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