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Haplogroup R1a
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| Haplogroup R1a | |
|---|---|
| Possible time of origin | 22,000[1] to 25,000[2] years ago |
| Possible place of origin | Eurasia |
| Ancestor | Haplogroup R1 |
| Descendants | R-M459, R-YP4141 |
| Defining mutations |
|
| Highest frequencies | See List of R1a frequency by population |

Haplogroup R1a (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 Central Asia, southern Siberia and South Asia.[3][2]
The R1a (R-M420) subclade diverged from R1 (R-M173) 15-25,000[2][4][5] years ago, its subclade M417 (R1a1a1) diversified c. 3,400-5,800 years ago.[6][5] 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
[edit]R1a origins
[edit]The genetic divergence of R1a (M420) is estimated to have occurred 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, c. 13,000 years ago),[7][8] and the earliest case of R* among Upper Paleolithic Ancient North Eurasians,[9] from which the Eastern Hunter-Gatherers predominantly derive their ancestry.[10] The genome of an individual belonging to the R1a5 subclade, dated to 10785–10626 BCE, from Peschanitsa, Arkhangelsk, Russia, and identified as a Western Russian Hunter-Gatherer, was published in January 2021.[11]
Diversification of R1a1a1 (M417) and ancient migrations
[edit]
According to Underhill et al. (2014), the downstream M417 (R1a1a1) subclade diversified into Z282 (R1a1a1b1a) and Z93 (R1a1a1b2) circa 5,800 years ago "in the vicinity of Iran and Eastern Turkey".[6][note 1] Even though R1a occurs as a Y-chromosome haplogroup among speakers of 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 Scandinavia[13][3] being most prevalent in Eastern Europe, Central 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.[14]
Indo-European relation
[edit]Proposed steppe dispersal of R1a1a
[edit]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.[15] 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.[16] 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.[17]
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.[18][19][20]
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 Indian subcontinent, consistent with multiple waves of arrival."[21] 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."[22][23]
Possible Yamnaya or Corded Ware origins
[edit]

David Anthony considers the Yamnaya culture to be the Indo-European Urheimat.[24][25] 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".[26] Yet, all their seven Yamnaya samples belonged to the R1b-M269 subclade,[26] 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.[27]
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.[28]
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.[29]
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.[30]
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.[31][note 2]
Proposed South Asian origins
[edit]Kivisild et al. (2003) have proposed either South or West Asia,[32][note 3] while Mirabal et al. (2009) see support for both South and Central Asia.[13] Sengupta et al. (2006) have proposed Indian origins.[33] Thanseem et al. (2006) have proposed either South or Central Asia.[34] Sahoo et al. (2006) have proposed either South or West Asia.[35] Thangaraj et al. (2010) have also proposed a South Asian origin.[36] Sharma et al.(2009) theorizes the existence of R1a in India beyond 18,000 years to possibly 44,000 years in origin.[1]
A number of studies from 2006 to 2010 concluded that South Asian populations have the highest STR diversity within R1a1a,[37][38][13][3][1][39] and subsequent older TMRCA datings.[note 4] R1a1a is present among both higher (Brahmin) castes and lower castes, and while the frequency is higher among Brahmin castes, the oldest TMRCA datings of the R1a haplogroup occur in the Saharia tribe, a scheduled caste of the Bundelkhand region of Central India.[1][39]
From these findings some researchers argued that R1a1a originated in South Asia,[38][1][note 5] excluding a more recent, yet minor, genetic influx from Indo-European migrants in northwestern regions such as Afghanistan, Balochistan, Punjab, and Kashmir.[38][37][3][note 6]
The conclusion that R1a originated in India has been questioned by more recent research,[21][41][note 7] offering an argument that R1a arrived in India with multiple waves of migration.[21][42]
Proposed Transcaucasia and West Asian origins and possible influence on Indus Valley Civilization
[edit]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 8] 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 9]
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[44] and expanded there during the Indus Valley civilization.[2][45]
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",[44] 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."[44] Yet, Lazaridis noted that sample I1635 of Lazaridis et al. (2016), their Armenian Kura-Araxes sample, carried Y-haplogroup R1b1-M415(xM269)[note 10] (also called R1b1a1b-CTS3187).[46][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 11]) may reflect this."[2][note 12] 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."[45][note 13]
However, according to Narasimhan et al. (2018), steppe pastoralists are a likely source for R1a in India.[48][note 14]
Phylogeny
[edit]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
[edit]The topology of R1a is as follows (codes [in brackets] non-isogg codes):[12][49][verification needed][50][2][51] 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."[52]
- P P295/PF5866/S8 (also known as K2b2).
- R (R-M207)[50][12]
- R*
- R1 (R-M173)
- R1*[50]
- R1a (M420)[50] (Eastern Europe, Asia)[2]
- R1a*[12]
- R1a1[50] (M459/PF6235,[50] SRY1532.2/SRY10831.2[50])
- R1a1 (M459)[50][12]
- R1a1a (M17, M198)[50]
- R1a1a1 (M417, page7)[50]
- R1a1a1a (CTS7083/L664/S298)[50]
- R1a1a1b (S224/Z645, S441/Z647)[50]
- R1a1a1b1 (PF6217/S339/Z283)[50]
- R1a1a1b1a (Z282)[50] [R1a1a1a*] (Z282)[53] (Eastern Europe)
- R1a1a1b1a1[50] [The old topological code is R1a1a1b*,which is outdated and might lead to some confusion.][53] (M458)[50][53] [R1a1a1g] (M458)[51]
- R1a1a1b1a2[50] (S466/Z280, S204/Z91)[50]
- R1a1a1b1a2a[50]
- R1a1a1b1a2b (CTS1211)[50] [R1a1a1c*] (M558)[53] [R-CTS1211] (V2803/CTS3607/S3363/M558, CTS1211/S3357, Y34/FGC36457)[12]
- R1a1a1b1a2b3* (M417+, Z645+, Z283+, Z282+, Z280+, CTS1211+, CTS3402, Y33+, CTS3318+, Y2613+) (Gwozdz's Cluster K)[49][verification needed]
- R1a1a1b1a2b3a (L365/S468)[50]
- R1a1a1b1a3 (Z284)[50] [R1a1a1a1] (Z284)[53]
- R1a1a1b1a (Z282)[50] [R1a1a1a*] (Z282)[53] (Eastern Europe)
- R1a1a1b2 (F992/S202/Z93)[50] [R1a1a2*] (Z93, M746)[53] (Central Asia, South Asia and West Asia)
- R1a1a1b1 (PF6217/S339/Z283)[50]
- [R1a1a1c] (M64.2, M87, M204)[51]
- [R1a1a1d] (P98)[51]
- [R1a1a1d2a][54]
- [R1a1a1e] (PK5)[51]
- R1a1a1 (M417, page7)[50]
- R1b (M343) (Western Europe)
- R2 (India)
Haplogroup R
[edit]| Haplogroup R phylogeny |
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]R1a, defined by the mutation M420, has two primary branches: R-M459 (R1a1) and R-YP4141 (R1a2).
As of 2025, ten ancient basal R1a* genotypes have been recovered and published, from remains found in Estonia, Poland, Russia, and Ukraine; the oldest sample (Vasilevka 497) dated to c. 8700 BCE, and excavated in the Vasylivka, Bakhmut Raion, Donetsk Oblast.[55][5]
R-YP4141 (R1a2)
[edit]R1a2 (R-YP4141) has two branches R1a2a (R-YP5018) and R1a2b (R-YP4132).[56]
This rare primary subclade was initially regarded as part of a paragroup of R1a*, defined by SRY1532.2 (and understood to always exclude M459 and its synonyms SRY10831.2, M448, L122, and M516).[3][57]
YP4141 later replaced SRY1532.2 – which was found to be unreliable – and the R1a(xR-M459) group was redefined as R1a2. It is relatively unusual, though it has been tested in more than one survey. Sahoo et al. (2006) reported R-SRY1532.2* for 1/15 Himachal Pradesh Rajput samples.[38] Underhill et al. (2009) reported 1/51 in Norway, 3/305 in Sweden, 1/57 Greek Macedonians, 1/150 (or 2/150) Iranians, 2/734 ethnic Armenians, 1/141 Kabardians, 1/121 Omanis, 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]
The oldest known example genotyped is from a set of remains, dating to c. 3500 BCE, recovered from the Kumyshanskaya Cave, in Russia.[5]
R-M459 (R1a1)
[edit]The major subclade R-M459 includes an overwhelming majority of individuals within R1a more broadly.
Ancient R-M459 genotypes, dating to c. 8650 BCE, have been recovered from two sets of remains excavated at Minino, Russia.[5]
R-YP1272 (R1a1b)
[edit]R-YP1272, also known as R-M459(xM198), is an extremely rare primary subclade of R1a1. It has been found in three individuals, from Belarus, Tunisia and the Coptic community in Egypt respectively.[58]
R-M17/M198 (R1a1a)
[edit]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)
[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]).
The oldest known basal R1a1a1 genotype so far published has been dated to c. 5650 BCE, and was recovered from a site at Trestiana, Romania.[5]
R-Z282 (R1a1a1b1a) (Eastern Europe)
[edit]This large subclade appears to encompass most of the R1a1a found in Europe.[17]
- 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.[59]
R-M458 (R1a1a1b1a1)
[edit]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." (Zhivotovsky speeds, 3x overvalued)[3] M458 was found in one skeleton from a 14th-century grave field in Usedom, Mecklenburg-Vorpommern, Germany.[60] The paper by Underhill et al. (2009) also reports a surprisingly high frequency of M458 in some Northern Caucasian populations (18% among Ak Nogai,[61] 7.8% among Qara Nogai and 3.4% among Abazas).[62]
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).[63] It apparently accounts for about 8% of Polish men, making it the most common subclade in Poland. Outside of Poland it is less common.[64] 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.[65]
R-M334
[edit]R-M334 ([R1a1a1g1],[51] a subclade of [R1a1a1g] (M458)[51] c.q. R1a1a1b1a1 (M458)[50]) 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)[49][verification needed] is a STR based group that is R-M17(xM458). This cluster is common in Poland but not exclusive to Poland.[65]
R1a1a1b1a2b3a (R-L365)
[edit]R1a1a1b1a2b3a (R-L365)[50] was early called Cluster G.[citation needed]
R1a1a1b2 (R-Z93) (Asia)
[edit]| 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).[17]
- 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] The oldest published R-Z93 genotypes being sampled from graves, dated to c. 2650 - 2700 BCE, in Naumovskoye, and Khanevo, Vologda Oblast, and Khaldeevo, Rostov District, Russia.[5]
- 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-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 (R1a1b2a2*) occurs at high frequency in South Asia: India, Pakistan, Afghanistan, and the Himalayas. Turkey share R1a (12.1%) sublineages.[66] Roma from Slovakia share 3% of R1a[67] 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
[edit]
Pre-historical
[edit]In Mesolithic Europe, R1a is characteristic of Eastern Hunter-Gatherers (EHGs).[68] 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.[69][70][68] A male, named PES001, from Peschanitsa in northwestern Russia was found to carry R1a5, and dates to at least 10,600 years ago.[7] More examples include the males Minino II (V) and Minino II (I/1), with the former carrying R1a1 and the latter R1a respectively, with the former being at 10,600 years old and the latter at least 10,400 years old respectively, both from Minino in northwestern Russia.[71] A Mesolithic male from Karelia c. 8,800 BCE to 7950 BCE has been found to be carrying haplogroup R1a.[72] A Mesolithic male buried at Deriivka c. 7000 BCE to 6700 BCE carried the paternal haplogroup R1a and the maternal U5a2a.[20] Another male from Karelia from c. 5,500 to 5,000 BC, who was considered an EHG, carried haplogroup R1a.[18] 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.[73] According to archaeologist David Anthony, the paternal R1a-Z93 was found at the Oskol river near a no longer existing kolkhoz "Alexandria", Ukraine c. 4000 BCE, "the earliest known sample to show the genetic adaptation to lactase persistence (13910-T)."[74] R1a has been found in the Corded Ware culture,[75][76] in which it is predominant.[77] Examined males of the Bronze Age Fatyanovo culture belong entirely to R1a, specifically subclade R1a-Z93.[68][69][78]
Haplogroup R1a has later been found in ancient fossils associated with the Urnfield culture;[79] as well as the burial of the remains of the Sintashta,[19] Andronovo,[80] the Pazyryk,[81] Tagar,[80] Tashtyk,[80] and Srubnaya cultures, the inhabitants of ancient Tanais,[82] in the Tarim mummies,[83] and the aristocracy of Xiongnu.[84] 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.[75]
Europe
[edit]In Europe, the R1a1a sub-clade is primarily characteristic of Balto-Slavic populations, with two exceptions: southern Slavs and northern Russians.[85] The highest frequency of R1a1a in Europe is observed in Sorbs (63%),[86] a West Slavic ethnic group, followed by Hungarians (60%).[15] Other groups with significant R1a1a, ranging from 27% to up to 58%, include Czechs, Poles, Slovenians, Slovaks, Moldovans, Belarusians, Rusyns, Ukrainians, and Russians.[85][86][15] R1a frequency decreases in northeastern Russian populations down to 20%–30%, in contrast to central-southern Russia, where its frequency is twice as high. In the Baltics, R1a1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).[87][88][89][15][90]
There is also a significant presence in peoples of Germanic descent, with highest levels in Norway, Sweden and Iceland, where between 20 and 30% of men are in R1a1a.[91][92] 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.[93][94] Haplogroup R1a1a averages between 10 and 30% in Germans, with a peak in Rostock at 31.3%.[95] R1a1a is found at a very low frequency among Dutch people (3.7%)[15] and is virtually absent in Danes.[96]
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.[97][better source needed] The Balkans shows wide variation between areas with significant levels of R1a1a, for example 36–39% in Slovenia,[98] 27–34% in Croatia,[88][99][100][101][102] and over 30% in Greek Macedonia, but less than 10% in Albania, Kosovo and parts of Greece south of Olympus gorge.[103][89][15]
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 Turkey, 12,1%[66][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.[104][105] 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 and West 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 15]
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.[106]
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.[107]
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).[108]
Malyarchuk et al. (2013) found R1a1-SRY10831.2 in 30.0% (12/40) of a sample of Tajiks from Tajikistan.[109]
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.[110]
South Asia
[edit]In South Asia, R1a1a has often been observed in a number of demographic groups.[38][37]
South Asian populations have the highest STR diversity within R1a1a,[37][38][13][3][1][39] and subsequent older TMRCA datings.[note 16] In India, high frequencies of this haplogroup is observed in West Bengal Brahmins (72%) in the east,[37] Bhanushali (67%) and Gujarat Lohanas (60%) in the west,[3] Uttar Pradesh Brahmins (68%), Punjab/Haryana Khatris (67%) and Ahirs (63%) in the north,[1][37][3] and Karnataka Medars (39%) in the south.[111] It has also been found in several South Indian Dravidian-speaking Adivasis including the Chenchu (26%) of Andhra Pradesh and Kota of Andhra Pradesh (22.58%)[112] and the Kallar of Tamil Nadu suggesting that R1a1a is widespread in Tribal Southern Indians.[32]
Besides these, studies show high percentages in regionally diverse groups such as Manipuris (50%)[3] to the extreme North East and among Punjabis (47%)[32] to the extreme North West.
In Pakistan it is found at 80% among Yusufzai tribe of Pashtuns (51%) from Swat District,[113] 71% among the Mohanna community 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.[114] Hindus of Chitwan District in the Terai region Nepal show it at 69%.[115]
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.[16][116][117]
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.[118] 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 Tibet (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. A different subclade of R1 was also found in 1.5% of a sample of northern Hui from Ningxia.[119] 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.[120] 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.[121]
In Eastern Siberia, R1a1a is found among certain indigenous ethnic groups including Kamchatkans and Chukotkans, and peaking in Itel'man at 22%.[122]
Southeast Asia
[edit]Y-haplogroups R1a-M420 and R2-M479 are found in Ede (8.3% and 4.2%) and Giarai (3.7% and 3.7%) peoples in Vietnam. The Cham additionally have haplogroups R-M17 (13.6%) and R-M124 (3.4%).
R1a1a1b2a2a (R-Z2123) and R1a1 are found in Khmer peoples from Thailand (3.4%) and Cambodia (7.2%) respectively. Haplogroup R1a1a1b2a1b (R-Y6) is also found among Kuy peoples (5%).
According to Changmai et al. (2022), these haplogroup frequencies originate from South Asians, who left a cultural and genetic legacy in Southeast Asia since the first millennium CE.[123]
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%.[124][125][126]
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.[127]
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).[108]
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.[109]
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
[edit]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.
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See also
[edit]Notes
[edit]- ^ According to Family Tree,[who?] they diversified c. 5,000 years ago.[12]
- ^ Semenov & 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 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, November 13–15, 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; 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.
- ^ 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."[32]
- ^ Sengupta (2006): "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."
- ^ South-Asian origins:
* 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.[40]
* 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." - ^ Though Sengupta (2006) did concede that "[R1a1 and R2] could have actually arrived in southern India from a southwestern Asian source region multiple times." In full: "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."[37]
- ^ Lalueza-Fox: "Some years ago, local scientists supported the view that the existence of an R1a Y chromosome was not attributable to a foreign gene flow but instead that this lineage had emerged on the subcontinent and spread from there. But the phylogenetic reconstruction of this haplogroup did not support this view."[41]
- ^ Yet, Haak et al. also explicitly state: "a type of Near Eastern ancestry different from that which was introduced by early farmers".[clarification needed][43]
- ^ According to Family Tree DNA, L664 formed 4,700 ybp, that is, 2,700 BCE.[12]
- ^ 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)[47]
- ^ According to Family Tree DNA, M780 formed 4700 ybp.[12] 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)."[48]
- ^ 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.
- ^ Sengupta (2006): "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."
References
[edit]- ^ a b c d e f g 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 Underhill et al. 2009.
- ^ "YTree v13.01.00 - R1". YFull.Com.
- ^ a b c d e f g "Welcome to FamilyTreeDNA Discover". FamilyTreeDNA Discover. Retrieved January 19, 2025.
- ^ a b Underhill et al. 2014, p. 130.
- ^ a b 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 Advances. 7 (4) eabd6535. Bibcode:2021SciA....7.6535S. doi:10.1126/sciadv.abd6535. PMC 7817100. PMID 33523926.
- ^ Haak, Wolfgang; Lazaridis, Iosif; Patterson, Nick; Rohland, Nadin; Mallick, Swapan; Llamas, Bastien; Brandt, Guido; Nordenfelt, Susanne; Harney, Eadaoin; Stewardson, Kristin; Fu, Qiaomei (February 10, 2015). "Massive migration from the steppe is a source for Indo-European languages in Europe". bioRxiv 013433. arXiv:1502.02783. doi:10.1101/013433. S2CID 196643946. Archived from the original on December 23, 2019. Retrieved February 8, 2021.
- ^ Raghavan, Maanasa; Skoglund, Pontus; Graf, Kelly E.; Metspalu, Mait; Albrechtsen, Anders; Moltke, Ida; Rasmussen, Simon; Stafford Jr, Thomas W.; Orlando, Ludovic; Metspalu, Ene; Karmin, Monika (January 2014). "Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans". Nature. 505 (7481): 87–91. Bibcode:2014Natur.505...87R. doi:10.1038/nature12736. PMC 4105016. PMID 24256729.
- ^ Narasimhan, Vagheesh M.; Patterson, Nick; Moorjani, Priya; Rohland, Nadin; Bernardos, Rebecca; Mallick, Swapan; Lazaridis, Iosif; Nakatsuka, Nathan; Olalde, Iñigo; Lipson, Mark; Kim, Alexander M. (September 6, 2019). "The formation of human populations in South and Central Asia". Science. 365 (6457) eaat7487. doi:10.1126/science.aat7487. PMC 6822619. PMID 31488661.
Y chromosome haplogroup types R1b or R1a not represented in Iran and Turan in this period ...
- ^ 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; Reidla, Maere; Rootsi, Siiri; Kivisild, Toomas; Scheib, Christiana Lyn (January 20, 2021). "Genetic ancestry changes in Stone to Bronze Age transition in the East European plain". Science Advances. 7 (4) eabd6535. doi:10.1126/sciadv.abd6535. PMC 7817100. PMID 33523926.
- ^ a b c d e f g h i j "R1a tree". YFull. Archived from the original on August 19, 2016. Retrieved July 15, 2016.
- ^ a b c d Mirabal et al. 2009.
- ^ Zerjal, T.; et al. (1999). "The use of Y-chromosomal DNA variation to investigate population history: recent male spread in Asia and Europe". In Papiha, S. S.; Deka, R. & Chakraborty, R. (eds.). Genomic diversity: applications in human population genetics. New York: Kluwer Academic/Plenum Publishers. pp. 91–101. ISBN 978-0-3064-6295-5.
- ^ a b c d e f Semino et al. 2000.
- ^ a b Wells 2001.
- ^ a b c Pamjav et al. 2012.
- ^ a b Haak et al. 2015.
- ^ a b Allentoft et al. 2015.
- ^ a b Mathieson et al. 2015.
- ^ a b c Silva et al. 2017.
- ^ Joseph, Tony (June 16, 2017). "How genetics is settling the Aryan migration debate". The Hindu. Archived from the original on October 4, 2023. Retrieved June 2, 2019.
- ^ Silva, Marina; Oliveira, Marisa; Vieira, Daniel; Brandão, Andreia; Rito, Teresa; Pereira, Joana B.; Fraser, Ross M.; Hudson, Bob; Gandini, Francesca; Edwards, Ceiridwen; Pala, Maria; Koch, John; Wilson, James F.; Pereira, Luísa; Richards, Martin B. (March 23, 2017). "A genetic chronology for the Indian Subcontinent points to heavily sex-biased dispersals". BMC Evolutionary Biology. 17 (1): 88. Bibcode:2017BMCEE..17...88S. doi:10.1186/s12862-017-0936-9. ISSN 1471-2148. PMC 5364613. PMID 28335724.
- ^ Anthony 2007.
- ^ Anthony & Ringe 2015.
- ^ a b Haak et al. 2015, p. 5.
- ^ Semenov & Bulat 2016.
- ^ Haber et al. 2012"R1a1a7-M458 was absent in Afghanistan, suggesting that R1a1a-M17 does not support, as previously thought [47], expansions from the Pontic Steppe [3], bringing the Indo-European languages to Central Asia and India."
- ^ Klejn, Leo S. (April 22, 2017). "The Steppe Hypothesis of Indo-European Origins Remains to be Proven". Acta Archaeologica. 88 (1): 193–204. doi:10.1111/j.1600-0390.2017.12184.x. ISSN 0065-101X. Archived from the original on December 25, 2022. Retrieved November 23, 2022. "As for the Y-chromosome, it was already noted in Haak, Lazaridis et al. (2015) that the Yamnaya from Samara had Y-chromosomes which belonged to R-M269 but did not belong to the clade common in Western Europe (p. 46 of supplement). Also, not a single R1a in Yamnaya unlike Corded Ware (R1a-dominated)."
- ^ Koch, John T.; Cunliffe, Barry (2016). Celtic from the West 3: Atlantic Europe in the Metal Ages. Oxbow Books. p. 634. ISBN 978-1-78570-228-0. Archived from the original on November 23, 2022. Retrieved November 23, 2022.
- ^ Semenov & Bulat 2016, p. 41.
- ^ a b c d Kivisild et al. 2003.
- ^ Sengupta S, Zhivotovsky LA, King R, Mehdi SQ, Edmonds CA, Chow CE, et al. (February 2006). "Polarity and temporality of high-resolution y-chromosome distributions in India identify both indigenous and exogenous expansions and reveal minor genetic influence of Central Asian pastoralists". American Journal of Human Genetics. 78 (2): 202–221. doi:10.1086/499411. PMC 1380230. PMID 16400607."Although considerable cultural impact on social hierarchy and language in South Asia is attributable to the arrival of nomadic Central Asian pastoralists, genetic data (mitochondrial and Y chromosomal) have yielded dramatically conflicting inferences on the genetic origins of tribes and castes of South Asia. We sought to resolve this conflict, using high-resolution data on 69 informative Y-chromosome binary markers and 10 microsatellite markers from a large set of geographically, socially, and linguistically representative ethnic groups of South Asia. 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. R1a1 and R2 haplogroups indicate demographic complexity that is inconsistent with a recent single history.ASSOCIATED MICROSATELLITE ANALYSES OF THE HIGH-FREQUENCY R1A1 HAPLOGROUP CHROMOSOMES INDICATE INDEPENDENT RECENT HISTORIES OF THE INDUS VALLEY AND THE PENINSULAR INDIAN REGION."
- ^ Thanseem I, Thangaraj K, Chaubey G, Singh VK, Bhaskar LV, Reddy BM, et al. (August 2006). "Genetic affinities among the lower castes and tribal groups of India: inference from Y chromosome and mitochondrial DNA". BMC Genetics. 7: 42. doi:10.1186/1471-2156-7-42. PMC 1569435. PMID 16893451.
- ^ Sahoo S, Singh A, Himabindu G, Banerjee J, Sitalaximi T, Gaikwad S, et al. (January 2006). "A prehistory of Indian Y chromosomes: evaluating demic diffusion scenarios". Proceedings of the National Academy of Sciences of the United States of America. 103 (4): 843–848. Bibcode:2006PNAS..103..843S. doi:10.1073/pnas.0507714103. PMC 1347984. PMID 16415161.
- ^ Thangaraj K, Naidu BP, Crivellaro F, Tamang R, Upadhyay S, Sharma VK, et al. (December 2010). Cordaux R (ed.). "The influence of natural barriers in shaping the genetic structure of Maharashtra populations". PLOS ONE. 5 (12) e15283. Bibcode:2010PLoSO...515283T. doi:10.1371/journal.pone.0015283. PMC 3004917. PMID 21187967.
- ^ a b c d e f g Sengupta 2006.
- ^ a b c d e f Sahoo et al. 2006.
- ^ a b c Thangaraj et al. 2010.
- ^ Sahoo et al. 2006, p. 845-846.
- ^ a b Lalueza-Fox, C. (2022). Inequality: A Genetic History. MIT Press. pp. 81–82. ISBN 978-0-262-04678-7. Archived from the original on July 16, 2023. Retrieved July 16, 2023.
- ^ Narasimhan et al. 2019.
- ^ Haak et al. 2015, p. 4.
- ^ a b c Mascarenhas et al. 2015, p. 9.
- ^ a b Poznik et al. 2016, p. 5.
- ^ Arame's English blog, Y DNA from ancient Near East Archived November 27, 2016, at the Wayback Machine
- ^ "Dienekes' Anthropology Blog: Major new article on the deep origins of Y-haplogroup R1a (Underhill et al. 2014)". March 27, 2014. Archived from the original on December 20, 2019. Retrieved December 20, 2019.[unreliable source?]
- ^ a b Narasimhan et al. 2018.
- ^ a b c "About Us". Family Tree DNA. Archived from the original on August 15, 2019. Retrieved December 20, 2019.
- ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa "ISOGG 2017 Y-DNA Haplogroup R". isogg.org. Archived from the original on February 10, 2007. Retrieved December 20, 2019.
- ^ a b c d e f g h i j k "Haplogroup R (Y-DNA) - SNPedia". www.snpedia.com. Archived from the original on May 5, 2018. Retrieved December 20, 2019.
- ^ Karafet et al. 2014.
- ^ a b c d e f g h i j k l m Underhill et al. 2014, p. 125.
- ^ "R1a in Yamnaya". Eurogenes Blog. March 21, 2016. Archived from the original on May 5, 2018. Retrieved December 20, 2019.
- ^ "haplotree.info - ancientdna.info. Map based on All Ancient DNA v. 2.07.26". haplotree.info. Retrieved January 19, 2025.
- ^ "R1a YTree".
- ^ Krahn, Thomas. "Draft Y-Chromosome Tree". Family Tree DNA. Archived from the original on May 26, 2013. Retrieved December 7, 2012.
- ^ "R-M459 YTree".
- ^ Regueiro 2006.
- ^ Freder, Janine (2010). Die mittelalterlichen Skelette von Usedom: Anthropologische Bearbeitung unter besonderer Berücksichtigung des ethnischen Hintergrundes [Anthropological investigation in due consideration of the ethnical background] (Thesis) (in German). Freie Universität Berlin. p. 86. doi:10.17169/refubium-8995.
- ^ https://cyberleninka.ru/article/n/tyurki-kavkaza-sravnitelnyy-analiz-genofondov-po-dannym-o-y-hromosome Archived November 7, 2023, at the Wayback Machine "высокая частота R1a среди кубанских ногайцев (субветвь R1a1a1g-M458 забирает 18%"
- ^ Underhill, P. A.; et al. (2009). "Separating the post-Glacial coancestry of European and Asian y chromosomes within haplogroup R1a". European Journal of Human Genetics. 18 (4): 479–484. doi:10.1038/ejhg.2009.194. PMC 2987245. PMID 19888303.
- ^ Gwozdz, Peter (August 6, 2018). "Polish Y-DNA Clades". Archived from the original on July 15, 2016. Retrieved July 15, 2016.
- ^ Pawlowski et al. 2002.
- ^ a b Gwozdz 2009.
- ^ a b Kars, M. E.; Başak, A. N.; Onat, O. E.; Bilguvar, K.; Choi, J.; Itan, Y.; Çağlar, C.; Palvadeau, R.; Casanova, J. L.; Cooper, D. N.; Stenson, P. D.; Yavuz, A.; Buluş, H.; Günel, M.; Friedman, J. M.; Özçelik, T. (2021). "The genetic structure of the Turkish population reveals high levels of variation and admixture". Proceedings of the National Academy of Sciences of the United States of America. 118 (36) e2026076118. Bibcode:2021PNAS..11826076K. doi:10.1073/pnas.2026076118. PMC 8433500. PMID 34426522.
- ^ Petrejcíková, EVA; Soták, Miroslav; Bernasovská, Jarmila; Bernasovský, Ivan; Sovicová, Adriana; Bôziková, Alexandra; Boronová, Iveta; Švícková, Petra; Gabriková, Dana; MacEková, Sona (2009). "Y-haplogroup frequencies in the Slovak Romany population". Anthropological Science. 117 (2): 89–94. doi:10.1537/ase.080422.
- ^ a b c Saag et al. 2020, p. 5.
- ^ a b Saag et al. 2020, p. 29, Table 1.
- ^ Saag et al. 2020, Supplementary Data 2, Row 4.
- ^ Posth, Cosimo; Yu, He; Ghalichi, Ayshin; Rougier, Hélène; Crevecoeur, Isabelle; Huang, Yilei; Ringbauer, Harald; Rohrlach, Adam B.; Nägele, Kathrin; Villalba-Mouco, Vanessa; Radzeviciute, Rita; Ferraz, Tiago; Stoessel, Alexander; Tukhbatova, Rezeda; Drucker, Dorothée G. (March 1, 2023). "Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers". Nature. 615 (7950): 117–126. Bibcode:2023Natur.615..117P. doi:10.1038/s41586-023-05726-0. hdl:10256/23099. ISSN 1476-4687. PMC 9977688. PMID 36859578.
- ^ Fu et al. 2016.
- ^ Saag et al. 2017.
- ^ Anthony 2019, pp. 16, 17.
- ^ a b Haak et al. 2008.
- ^ Brandit et al. 2013.
- ^ Malmström et al. 2019, p. 2.
- ^ Saag et al. 2020, Supplementary Data 2, Rows 5-49.
- ^ Schweitzer, D. (March 23, 2008). "Lichtenstein Cave Data Analysis" (PDF). dirkschweitzer.net. Archived from the original (PDF) on August 14, 2011. Summary in English of Schilz (2006).
- ^ a b c Keyser et al. 2009.
- ^ Ricaut et al. 2004.
- ^ Korniyenko, I. V.; Vodolazhsky D. I. "Использование нерекомбинантных маркеров Y-хромосомы в исследованиях древних популяций (на примере поселения Танаис)" [The use of non-recombinant markers of the Y-chromosome in the study of ancient populations (on the example of the settlement of Tanais)]. Материалы Донских антропологических чтений [Materials of the Don Anthropological Readings]. Rostov-on-Don: Rostov Research Institute of Oncology, 2013.
- ^ Chunxiang Li et al. 2010.
- ^ Kim et al. 2010.
- ^ a b Balanovsky et al. 2008.
- ^ a b Behar et al. 2003.
- ^ Kasperaviciūte, Kucinskas & Stoneking 2005.
- ^ a b Battaglia et al. 2008.
- ^ a b Rosser et al. 2000.
- ^ Tambets et al. 2004.
- ^ Bowden et al. 2008.
- ^ Dupuy et al. 2005.
- ^ Passarino et al. 2002.
- ^ Capelli et al. 2003.
- ^ Kayser et al. 2005.
- ^ Sanchez, J; Børsting, C; Hallenberg, C; Buchard, A; Hernandez, A; Morling, N (2003). "Multiplex PCR and minisequencing of SNPs—a model with 35 Y chromosome SNPs". Forensic Science International. 137 (1): 74–84. doi:10.1016/S0379-0738(03)00299-8. PMID 14550618.
- ^ Scozzari et al. 2001.
- ^ Underhill, Peter A. (January 1, 2015). "The phylogenetic and geographic structure of Y-chromosome haplogroup R1a". European Journal of Human Genetics. 23 (1): 124–131. doi:10.1038/ejhg.2014.50. PMC 4266736. PMID 24667786.
- ^ L. Barać; et al. (2003). "Y chromosomal heritage of Croatian population and its island isolates". European Journal of Human Genetics. 11 (7): 535–42. doi:10.1038/sj.ejhg.5200992. PMID 12825075. S2CID 15822710.
- ^ S. Rootsi; et al. (2004). "Phylogeography of Y-Chromosome Haplogroup I Reveals Distinct Domains of Prehistoric Gene Flow in Europe" (PDF). American Journal of Human Genetics. 75 (1): 128–137. doi:10.1086/422196. PMC 1181996. PMID 15162323. Archived from the original (PDF) on September 5, 2020. Retrieved February 13, 2021.
- ^ M. Peričić; et al. (2005). "High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations". Molecular Biology and Evolution. 22 (10): 1964–75. doi:10.1093/molbev/msi185. PMID 15944443.
- ^ M. Peričić; et al. (2005). "Review of Croatian Genetic Heritage as Revealed by Mitochondrial DNA and Y Chromosomal Lineages". Croatian Medical Journal. 46 (4): 502–513. PMID 16100752.
- ^ Pericić et al. 2005.
- ^ "Untitled". pereformat.ru (in Russian). Archived from the original on March 15, 2016. Retrieved May 29, 2017.
- ^ "Untitled". www.rodstvo.ru. Archived from the original on September 16, 2021. Retrieved May 29, 2017.
- ^ Zerjal et al. 2002.
- ^ Haber et al. 2012.
- ^ a b Di Cristofaro et al. 2013.
- ^ a b Malyarchuk et al. 2013.
- ^ Ashirbekov et al. 2017.
- ^ Shah 2011.
- ^ Arunkumar 2012.
- ^ Tariq, Muhammad; Ahmad, Habib; Hemphill, Brian E.; Farooq, Umar; Schurr, Theodore G. (2022). "Contrasting maternal and paternal genetic histories among five ethnic groups from Khyber Pakhtunkhwa, Pakistan". Scientific Reports. 12 (1): 1027. Bibcode:2022NatSR..12.1027T. doi:10.1038/s41598-022-05076-3. PMC 8770644. PMID 35046511.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Toomas Kivisild; Siiri Rootsi; Mait Metspalu; Ene Metspalu; Juri Parik; Katrin Kaldma; Esien Usanga; Sarabjit Mastana; Surinder S. Papiha; Richard Villems. "The Genetics of Language and Farming Spread in India" (PDF). In P. Bellwwood; C. Renfrew (eds.). Examining the farming/language dispersal hypothesis. McDonald Institute Monographs. Cambridge University. pp. 215–222. Archived from the original (PDF) on February 19, 2006. Retrieved December 20, 2019.
- ^ Fornarino et al. 2009.
- ^ Wang et al. 2003.
- ^ Zhou et al. 2007.
- ^ Liu Shu-hu et al. 2018.
- ^ Zhong et al. 2011.
- ^ Zhong, Hua; Shi, Hong; Qi, Xue-Bin; Duan, Zi-Yuan; Tan, Ping-Ping; Jin, Li; Su, Bing; Ma, Runlin Z. (2011). "Extended Y Chromosome Investigation Suggests Postglacial Migrations of Modern Humans into East Asia via the Northern Route". Molecular Biology and Evolution. 28 (1): 717–727. doi:10.1093/molbev/msq247. PMID 20837606.
- ^ Shou, Wei-Hua; Qiao, Wn-Fa; Wei, Chuan-Yu; Dong, Yong-Li; Tan, Si-Jie; Shi, Hong; Tang, Wen-Ru; Xiao, Chun-Jie (2010). "Y-chromosome distributions among populations in Northwest China identify significant contribution from Central Asian pastoralists and lesser influence of western Eurasians". Journal of Human Genetics. 55 (5): 314–322. doi:10.1038/jhg.2010.30. PMID 20414255. S2CID 23002493.
- ^ Lell et al. 2002.
- ^ Changmai, Piya; Jaisamut, Kitipong; Kampuansai, Jatupol; et al. (2022). "Indian genetic heritage in Southeast Asian populations". PLOS Genetics. 18 (2) e1010036. doi:10.1371/journal.pgen.1010036. PMC 8853555. PMID 35176016.
- ^ Mohammad et al. 2009.
- ^ Nasidze et al. 2004.
- ^ Nasidze et al. 2005.
- ^ Grugni et al. 2012.
Sources
[edit]- Allentoft, Morten E.; Sikora, Martin; Sjögren, Karl-Göran; Rasmussen, Simon; Rasmussen, Morten; Stenderup, Jesper; Damgaard, Peter B.; Schroeder, Hannes; et al. (2015). "Population genomics of Bronze Age Eurasia". Nature. 522 (7555): 167–172. Bibcode:2015Natur.522..167A. doi:10.1038/nature14507. PMID 26062507. S2CID 4399103. Archived from the original on July 13, 2019. Retrieved July 23, 2019.
- Anthony, David W. (2007), The Horse The Wheel And Language. How Bronze-Age Riders From the Eurasian Steppes Shaped The Modern World, Princeton University Press
- Anthony, David (Spring–Summer 2019). "Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard". Journal of Indo-European Studies. 47 (1–2). Archived from the original on May 3, 2024. Retrieved January 9, 2020.
- Anthony, David; Ringe, Don (2015), "The Indo-European Homeland from Linguistic and Archaeological Perspectives", Annual Review of Linguistics, 1: 199–219, doi:10.1146/annurev-linguist-030514-124812
- Shah, A. M.; Tamang, R.; Moorjani, P.; Rani, D. S.; Govindaraj, P.; Kulkarni, G.; Bhattacharya, T.; Mustak, M. S.; Bhaskar, L. V. K. S.; Reddy, A. G.; Gadhvi, D.; Gai, P. B.; Chaubey, G.; Patterson, N.; Reich, D.; Tyler-Smith, C.; Singh, L.; Thangaraj, K. (2011). "Indian Siddis: African Descendants with Indian Admixture". The American Journal of Human Genetics. 89 (1): 154–61. doi:10.1016/j.ajhg.2011.05.030. PMC 3135801. PMID 21741027.
- ArunKumar, G; Soria-Hernanz, DF; Kavitha, VJ; Arun, VS; Syama, A; Ashokan, KS (2012). "Population Differentiation of Southern Indian Male Lineages Correlates with Agricultural Expansions Predating the Caste System". PLOS ONE. 7 (11) e50269. Bibcode:2012PLoSO...750269A. doi:10.1371/journal.pone.0050269. PMC 3508930. PMID 23209694.
- Ashirbekov, E. E.; et al. (2017). "Distribution of Y-Chromosome Haplogroups of the Kazakh from the South Kazakhstan, Zhambyl, and Almaty Regions" (PDF). Reports of the National Academy of Sciences of the Republic of Kazakhstan. 6 (316): 85–95. Archived from the original (PDF) on September 23, 2021. Retrieved June 4, 2020.
- Balanovsky O, Rootsi S, Pshenichnov A, Kivisild T, Churnosov M, Evseeva I, Pocheshkhova E, Boldyreva M, et al. (2008). "Two Sources of the Russian Patrilineal Heritage in Their Eurasian Context". American Journal of Human Genetics. 82 (1): 236–250. doi:10.1016/j.ajhg.2007.09.019. PMC 2253976. PMID 18179905.
- Балановский, О. П. (November 30, 2015). Генофонд Европы (in Russian). KMK Scientific Press. ISBN 978-5-9907157-0-7. Archived from the original on May 3, 2024. Retrieved August 8, 2020.
- Battaglia V, Fornarino S, Al-Zahery N, Olivieri A, Pala M, Myres NM, King RJ, Rootsi S, et al. (2008). "Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe". European Journal of Human Genetics. 17 (6): 820–30. doi:10.1038/ejhg.2008.249. PMC 2947100. PMID 19107149.
- Behar D, Thomas MG, Skorecki K, Hammer MF, Bulygina E, Rosengarten D, Jones AL, Held K, et al. (2003). "Multiple Origins of Ashkenazi Levites: Y Chromosome Evidence for Both Near Eastern and European Ancestries" (PDF). American Journal of Human Genetics. 73 (4): 768–779. doi:10.1086/378506. PMC 1180600. PMID 13680527. Archived (PDF) from the original on April 17, 2007. Retrieved April 23, 2007.
- Bowden GR, Balaresque P, King TE, Hansen Z, Lee AC, Pergl-Wilson G, Hurley E, Roberts SJ, et al. (2008). "Excavating Past Population Structures by Surname-Based Sampling: The Genetic Legacy of the Vikings in Northwest England". Molecular Biology and Evolution. 25 (2): 301–309. doi:10.1093/molbev/msm255. PMC 2628767. PMID 18032405.
- Brandit, G.; et al. (The Genographic Consortium) (2013). "Ancient DNA Reveals Key Stages in the Formation of Central European Mitochondrial Genetic Diversity". Science. 342 (6155): 257–261. Bibcode:2013Sci...342..257B. doi:10.1126/science.1241844. PMC 4039305. PMID 24115443.
- Capelli C, Redhead N, Abernethy JK, Gratrix F, Wilson JF, Moen T, Hervig T, Richards M, et al. (2003). "A Y Chromosome Census of the British Isles" (PDF). Current Biology. 13 (11): 979–84. Bibcode:2003CBio...13..979C. doi:10.1016/S0960-9822(03)00373-7. PMID 12781138. S2CID 526263. Archived (PDF) from the original on July 8, 2021. Retrieved August 8, 2020. also at "University College London" (PDF). Archived (PDF) from the original on January 17, 2017. Retrieved January 29, 2007.
- Chunxiang Li; Hongjie Li; Yinqiu Cui; Chengzhi Xie; Dawei Cai; Wenying Li; Victor H Mair; Zhi Xu; et al. (2010). "Evidence that a West-East admixed population lived in the Tarim Basin as early as the early Bronze Age". BMC Biology. 8 (1): 15. doi:10.1186/1741-7007-8-15. PMC 2838831. PMID 20163704.
- Di Cristofaro J, Pennarun E, Mazières S, Myres NM, Lin AA, Temori SA, Metspalu M, Metspalu E, et al. (2013). "Afghan Hindu Kush: Where Eurasian Sub-Continent Gene Flows Converge". PLOS ONE. 8 (10). e76748. Bibcode:2013PLoSO...876748D. doi:10.1371/journal.pone.0076748. PMC 3799995. PMID 24204668.
- Dupuy BM, Stenersen M, Lu TT, Olaisen B (2005). "Geographical heterogeneity of Y-chromosomal lineages in Norway" (PDF). Forensic Science International. 164 (1): 10–19. doi:10.1016/j.forsciint.2005.11.009. PMID 16337760. Archived (PDF) from the original on July 22, 2012. Retrieved November 18, 2009.
- Fornarino, Simona; Pala, Maria; Battaglia, Vincenza; Maranta, Ramona; Achilli, Alessandro; Modiano, Guido; Torroni, Antonio; Semino, Ornella; et al. (2009). "Mitochondrial and Y-chromosome diversity of the Tharus (Nepal): a reservoir of genetic variation". BMC Evolutionary Biology. 9 (1): 154. Bibcode:2009BMCEE...9..154F. doi:10.1186/1471-2148-9-154. PMC 2720951. PMID 19573232.
- Fu, Qiaomei; et al. (May 2, 2016). "The genetic history of Ice Age Europe". Nature. 534 (7606): 200–205. Bibcode:2016Natur.534..200F. doi:10.1038/nature17993. hdl:10211.3/198594. PMC 4943878. PMID 27135931.
- Grugni V, Battaglia V, Kashani BH, Parolo S, Al-Zahery N, Achilli A, Olivieri A, Gandini F, Houshmand M, Sanati MH, Torroni A, Semino O (2012). "Ancient Migratory Events in the Middle East: New Clues from the Y-Chromosome Variation of Modern Iranians". PLOS ONE. 7 (7). e41252. Bibcode:2012PLoSO...741252G. doi:10.1371/journal.pone.0041252. PMC 3399854. PMID 22815981.
- Gwozdz (2009). "Y-STR Mountains in Haplospace, Part II: Application to Common Polish Clades" (PDF). Journal of Genetic Genealogy. 5 (2). Archived (PDF) from the original on July 21, 2011. Retrieved November 28, 2009.
- Haak, W.; Brandt, G.; Jong, H. N. d.; Meyer, C.; Ganslmeier, R.; Heyd, V.; Hawkesworth, C.; Pike, A. W. G.; et al. (2008). "Ancient DNA, Strontium isotopes, and osteological analyses shed light on social and kinship organization of the Later Stone Age". Proceedings of the National Academy of Sciences. 105 (47): 18226–18231. Bibcode:2008PNAS..10518226H. doi:10.1073/pnas.0807592105. PMC 2587582. PMID 19015520.
- Haak, Wolfgang; Lazaridis, Iosif; Patterson, Nick; Rohland, Nadin; Mallick, Swapan; Llamas, Bastien; Brandt, Guido; Nordenfelt, Susanne; et al. (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.
- Haber M, Platt DE, Ashrafian Bonab M, Youhanna SC, Soria-Hernanz DF, Martínez-Cruz B, Douaihy B, Ghassibe-Sabbagh M, et al. (2012). "Afghanistan's ethnic groups share a Y-chromosomal heritage structured by historical events". PLOS ONE. 7 (3). e34288. Bibcode:2012PLoSO...734288H. doi:10.1371/journal.pone.0034288. PMC 3314501. PMID 22470552.
- ISOGG (2012). "Y-DNA Haplogroup O and its Subclades - 2012".
- Karafet, Tatiana M.; Mendez, Fernando L.; Sudoyo, Herawati; Lansing, J. Stephen; Hammer, Michael F. (2014). "Improved phylogenetic resolution and rapid diversification of Y-chromosome haplogroup K-M526 in Southeast Asia". Nature. 23 (3): 369–373. doi:10.1038/ejhg.2014.106. PMC 4326703. PMID 24896152.
- Kasperaviciūte, D.; Kucinskas, V.; Stoneking, M. (2005). "Y Chromosome and Mitochondrial DNA Variation in Lithuanians". Annals of Human Genetics. 68 (5): 438–452. doi:10.1046/j.1529-8817.2003.00119.x. PMID 15469421. S2CID 26562505.
- Kayser M, Lao O, Anslinger K, Augustin C, Bargel G, Edelmann J, Elias S, Heinrich M, et al. (2005). "Significant genetic differentiation between Poland and Germany follows present-day political borders, as revealed by Y-chromosome analysis" (PDF). Human Genetics. 117 (5): 428–443. doi:10.1007/s00439-005-1333-9. PMID 15959808. S2CID 11066186. Archived from the original (PDF) on March 4, 2009.
- Keyser, Christine; Bouakaze, Caroline; Crubézy, Eric; Nikolaev, Valery G.; Montagnon, Daniel; Reis, Tatiana; Ludes, Bertrand (2009). "Ancient DNA provides new insights into the history of south Siberian Kurgan people". Human Genetics. 126 (3): 395–410. doi:10.1007/s00439-009-0683-0. PMID 19449030. S2CID 21347353.
- Kim, Kijeong; Brenner, Charles H.; Mair, Victor H.; Lee, Kwang-Ho; Kim, Jae-Hyun; Gelegdorj, Eregzen; Batbold, Natsag; Song, Yi-Chung; et al. (2010). "A western Eurasian male is found in 2000-year-old elite Xiongnu cemetery in Northeast Mongolia". American Journal of Physical Anthropology. 142 (3): 429–440. doi:10.1002/ajpa.21242. PMID 20091844.
- Kivisild, T; Rootsi, S; Metspalu, M; Mastana, S; Kaldma, K; Parik, J; Metspalu, E; Adojaan, M; et al. (2003). "The Genetic Heritage of the Earliest Settlers Persists Both in Indian Tribal and Caste Populations". AJHG. 72 (2): 313–32. doi:10.1086/346068. PMC 379225. PMID 12536373.
- Lazaridis, Iosif; et al. (2016). "Genomic insights into the origin of farming in the ancient Near East". Nature. 536 (7617): 419–424. Bibcode:2016Natur.536..419L. doi:10.1038/nature19310. PMC 5003663. PMID 27459054.
- Lell JT, Sukernik RI, Starikovskaya YB, Su B, Jin L, Schurr TG, Underhill PA, Wallace DC (2002). "The Dual Origin and Siberian Affinities of Native American Y Chromosomes" (PDF). American Journal of Human Genetics. 70 (1): 192–206. doi:10.1086/338457. PMC 384887. PMID 11731934. Archived from the original (PDF) on April 22, 2003.
- Liu Shu-hu; Nizam Yilihamu; Rabiyamu Bake; Abdukeram Bupatima; Dolkun Matyusup (2018). "A study of genetic diversity of three isolated populations in Xinjiang using Y-SNP". Acta Anthropologica Sinica. 37 (1): 146–156.
- Carlos Quiles (September 10, 2018). "A study of genetic diversity of three isolated populations in Xinjiang using Y-SNP". Indo-European.eu. Archived from the original on September 20, 2020. Retrieved July 24, 2020.
- Malmström, Helena; Günther, Torsten; Svensson, Emma M.; Juras, Anna; Fraser, Magdalena; Munters, Arielle R.; Pospieszny, Łukasz; Tõrv, Mari; et al. (October 9, 2019). "The genomic ancestry of the Scandinavian Battle Axe Culture people and their relation to the broader Corded Ware horizon". Proceedings of the Royal Society B. 286 (1912). doi:10.1098/rspb.2019.1528. PMC 6790770. PMID 31594508.
- Malyarchuk, Boris; Derenko, Miroslava; Wozniak, Marcin; Grzybowski, Tomasz (2013). "Y-chromosome variation in Tajiks and Iranians". Annals of Human Biology. 40 (1): 48–54. doi:10.3109/03014460.2012.747628. PMID 23198991. S2CID 2752490.
- Mascarenhas, Desmond D.; Raina, Anupuma; Aston, Christopher E.; Sanghera, Dharambir K. (2015). "Genetic and Cultural Reconstruction of the Migration of an Ancient Lineage". BioMed Research International. 2015 651415. doi:10.1155/2015/651415. PMC 4605215. PMID 26491681.
- Mathieson, Iain; Lazaridis, Iosif; Rohland, Nadin; Mallick, Swapan; Patterson, Nick; Alpaslan Roodenberg, Songul; Harney, Eadaoin; Stewardson, Kristin; et al. (2015). "Eight thousand years of natural selection in Europe". bioRxiv 10.1101/016477.
- Mirabal, Sheyla; Regueiro, M; Cadenas, AM; Cavalli-Sforza, LL; Underhill, PA; Verbenko, DA; Limborska, SA; Herrera, RJ; et al. (2009). "Y-Chromosome distribution within the geo-linguistic landscape of northwestern Russia". European Journal of Human Genetics. 17 (10): 1260–1273. doi:10.1038/ejhg.2009.6. PMC 2986641. PMID 19259129.
- Mohammad T, Xue Y, Evison M, Tyler-Smith C (2009). "Genetic structure of nomadic Bedouin from Kuwait". Heredity. 103 (5): 425–433. doi:10.1038/hdy.2009.72. PMC 2869035. PMID 19639002.
- Narasimhan, Vagheesh M.; Anthony, David; Mallory, James; Reich, David (2018). "The Genomic Formation of South and Central Asia". bioRxiv 10.1101/292581.
- Narasimhan, Vagheesh M.; Patterson, N.J.; Moorjani, Priya; Rohland, Nadin; et al. (2019), "The Formation of Human Populations in South and Central Asia", Science, 365 (6457) eaat7487, doi:10.1126/science.aat7487, PMC 6822619, PMID 31488661
- Nasidze I, Ling EY, Quinque D, Dupanloup I, Cordaux R, Rychkov S, Naumova O, Zhukova O, et al. (2004). "Mitochondrial DNA and Y-Chromosome Variation in the Caucasus" (PDF). Annals of Human Genetics. 68 (Pt 3): 205–221. doi:10.1046/j.1529-8817.2004.00092.x. PMID 15180701. S2CID 27204150. Archived from the original (PDF) on October 30, 2004.
- Nasidze I, Quinque D, Ozturk M, Bendukidze N, Stoneking M (2005). "MtDNA and Y-chromosome Variation in Kurdish Groups" (PDF). Annals of Human Genetics. 69 (Pt 4): 401–412. doi:10.1046/j.1529-8817.2005.00174.x. PMID 15996169. S2CID 23771698. Archived from the original (PDF) on August 23, 2009.
- Pamjav, Horolma; Fehér, Tibor; Németh, Endre; Pádár, Zsolt (2012), "Brief communication: new Y-chromosome binary markers improve phylogenetic resolution within haplogroup R1a1", American Journal of Physical Anthropology, 149 (4): 611–615, doi:10.1002/ajpa.22167, PMID 23115110, S2CID 4820868
- Passarino G, Cavalleri GL, Lin AA, Cavalli-Sforza LL, Børresen-Dale AL, Underhill (2002). "Different genetic components in the Norwegian population revealed by the analysis of mtDNA and Y chromosome polymorphisms". European Journal of Human Genetics. 10 (9): 521–529. doi:10.1038/sj.ejhg.5200834. PMID 12173029.
- Pathak, Ajai K.; Kadian, Anurag; Kushniarevich, Alena; Montinaro, Francesco; Mondal, Mayukh; Ongaro, Linda; Singh, Manvendra; Kumar, Pramod; et al. (December 6, 2018). "The Genetic Ancestry of Modern Indus Valley Populations from Northwest India". The American Journal of Human Genetics. 103 (6): 918–929. doi:10.1016/j.ajhg.2018.10.022. PMC 6288199. PMID 30526867.
- Pawlowski, R; Dettlaff-Kakol, A; MacIejewska, A; Paszkowska, R; Reichert, M; Jezierski, G (2002). "Population genetics of 9 Y-chromosome STR loci w Northern Poland". Arch. Med. Sadowej Kryminol. 52 (4): 261–277. PMID 14669672.
- Pericić M, Lauc LB, Klarić IM, Rootsi S, Janićijević B, Rudan I, Terzić R, Colak I, et al. (2005). "High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations". Mol. Biol. Evol. 22 (10): 1964–75. doi:10.1093/molbev/msi185. PMID 15944443.
- Poznik GD, et al. (2016). "Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences". Nature Genetics. 48 (6): 593–599. doi:10.1038/ng.3559. hdl:11858/00-001M-0000-002A-F024-C. PMC 4884158. PMID 27111036.
- Regueiro, M; Cadenas, AM; Gayden, T; Underhill, PA; Herrera, RJ (2006). "Iran: Tricontinental Nexus for Y-Chromosome Driven Migration". Hum Hered. 61 (3): 132–143. doi:10.1159/000093774. PMID 16770078. S2CID 7017701.
- Ricaut F, Keyser-Tracqui C, Bourgeois I, Crubézy E, Ludes B (2004). "Genetic Analysis of a Scytho-Siberian Skeleton and Its Implications for Ancient Central Asian Migrations". Human Biology. 76 (1): 109–25. doi:10.1353/hub.2004.0025. PMID 15222683. S2CID 35948291.
- Rosser ZH, Zerjal T, Hurles ME, Adojaan M, Alavantic D, Amorim A, Amos W, Armenteros M, et al. (2000). "Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language". American Journal of Human Genetics. 67 (6): 1526–1543. doi:10.1086/316890. PMC 1287948. PMID 11078479.
- Saag, Lehti; Varul, Liivi; Scheib, Christiana Lyn; Stenderup, Jesper; Allentoft, Morten E.; Saag, Lauri; Pagani, Luca; Reidla, Maere; et al. (July 24, 2017). "Extensive Farming in Estonia Started through a Sex-Biased Migration from the Steppe". Current Biology. 27 (14). Cell Press: 2185–2193. Bibcode:2017CBio...27E2185S. doi:10.1016/j.cub.2017.06.022. PMID 28712569.
- Saag, Lehti; Vasilyev, Sergey V.; Varul, Liivi; Kosorukova, Natalia V.; Gerasimov, Dmitri V.; Oshibkina, Svetlana V.; Griffith, Samuel J.; Solnik, Anu; et al. (July 3, 2020). "Genetic ancestry changes in Stone to Bronze Age transition in the East European plain". bioRxiv 10.1101/2020.07.02.184507.
- Sahoo, S; Singh, A; Himabindu, G; Banerjee, J; Sitalaximi, T; Gaikwad, S; Trivedi, R; Endicott, P; et al. (2006). "A prehistory of Indian Y chromosomes: Evaluating demic diffusion scenarios". Proceedings of the National Academy of Sciences. 103 (4): 843–848. Bibcode:2006PNAS..103..843S. doi:10.1073/pnas.0507714103. PMC 1347984. PMID 16415161.
- Scozzari R, Cruciani F, Pangrazio A, Santolamazza P, Vona G, Moral P, Latini V, Varesi L, et al. (2001). "Human Y-Chromosome Variation in the Western Mediterranean Area: Implications for the Peopling of the Region" (PDF). Human Immunology. 62 (9): 871–84. CiteSeerX 10.1.1.408.4857. doi:10.1016/S0198-8859(01)00286-5. PMID 11543889. Archived from the original (PDF) on December 17, 2012. Retrieved March 15, 2009.
- Semenov, Alexander S.; Bulat, Vladimir V. (2016), "Ancient Paleo-DNA of Pre-Copper Age North-Eastern Europe: Establishing the Migration Traces of R1a1 Y-DNA Haplogroup", European Journal of Molecular Biotechnology, 11 (1): 40–54, doi:10.13187/ejmb.2016.11.40, S2CID 172131289
- Semino, O; Passarino, G; Oefner, PJ; Lin, AA; Arbuzova, S; Beckman, LE; De Benedictis, G; Francalacci, P; et al. (2000). "The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective" (PDF). Science. 290 (5494): 1155–1159. Bibcode:2000Sci...290.1155S. doi:10.1126/science.290.5494.1155. PMID 11073453. Archived from the original (PDF) on November 25, 2003.
- Sengupta, S; Zhivotovsky, LA; King, R; Mehdi, SQ; Edmonds, CA; Chow, CE; Lin, AA; Mitra, M; et al. (2006). "Polarity and Temporality of High-Resolution Y-Chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influence of Central Asian Pastoralists". American Journal of Human Genetics. 78 (2): 202–21. doi:10.1086/499411. PMC 1380230. PMID 16400607.
- Sharma, S; Rai, E; Sharma, P; Jena, M; Singh, S; Darvishi, K; Bhat, AK; Bhanwer, AJ; et al. (2009). "The Indian origin of paternal haplogroup R1a1(*)substantiates the autochthonous origin of Brahmins and the caste system". Journal of Human Genetics. 54 (1): 47–55. doi:10.1038/jhg.2008.2. PMID 19158816.
- Schilz, Felix (2006). Molekulargenetische Verwandtschaftsanalysen am prähistorischen Skelettkollektiv der Lichtensteinhöhle [Molecular genetic kinship analysis on the prehistoric skeleton collective of the Lichtenstein Cave] (PDF) (Dissertation) (in German). Göttingen: Mathematisch-Naturwissenschaftlichen Fakultäten der Georg-August-Universität. Archived (PDF) from the original on June 8, 2011. Retrieved November 9, 2009.
- Silva, Marina; et al. (2017). "A genetic chronology for the Indian Subcontinent points to heavily sex-biased dispersals". BMC Evolutionary Biology. 17 (1): 88. Bibcode:2017BMCEE..17...88S. doi:10.1186/s12862-017-0936-9. PMC 5364613. PMID 28335724.
- Tambets K, Rootsi S, Kivisild T, Help H, Serk P, Loogväli EL, Tolk HV, Reidla M, et al. (2004). "The Western and Eastern Roots of the Saami—the Story of Genetic 'Outliers' Told by Mitochondrial DNA and Y Chromosomes". American Journal of Human Genetics. 74 (4): 661–682. doi:10.1086/383203. PMC 1181943. PMID 15024688.
- Thangaraj, Kumarasamy; Naidu, B. Prathap; Crivellaro, Federica; Tamang, Rakesh; Upadhyay, Shashank; Sharma, Varun Kumar; Reddy, Alla G.; Walimbe, S. R.; et al. (2010). Cordaux, Richard (ed.). "The Influence of Natural Barriers in Shaping the Genetic Structure of Maharashtra Populations". PLOS ONE. 5 (12) e15283. Bibcode:2010PLoSO...515283T. doi:10.1371/journal.pone.0015283. PMC 3004917. PMID 21187967.
- Underhill, PA; Myres, NM; Rootsi, S; Metspalu, M; Zhivotovsky, LA; King, RJ; Lin, AA; Chow, CE; et al. (November 4, 2009). "Separating the post-Glacial coancestry of European and Asian Y chromosomes within haplogroup R1a". European Journal of Human Genetics. 18 (4) (published April 2010): 479–84. doi:10.1038/ejhg.2009.194. PMC 2987245. PMID 19888303.
- Underhill, Peter A.; et al. (March 26, 2014). "The phylogenetic and geographic structure of Y-chromosome haplogroup R1a". European Journal of Human Genetics. 23 (1) (published January 2015): 124–131. doi:10.1038/ejhg.2014.50. PMC 4266736. PMID 24667786. "PDF" (PDF). Archived from the original (PDF) on August 16, 2016. Retrieved June 12, 2016.
- Wang, Wei; Wise, Cheryl; Baric, Tom; Black, Michael L.; Bittles, Alan H. (2003). "The origins and genetic structure of three co-resident Chinese Muslim populations: The Salar, Bo'an and Dongxiang". Human Genetics. 113 (3): 244–52. doi:10.1007/s00439-003-0948-y. PMID 12759817. S2CID 11138499.
- Wells, R.S. (2001), "The Eurasian Heartland: A continental perspective on Y-chromosome diversity", Proceedings of the National Academy of Sciences of the USA, 98 (18): 10244–10249, Bibcode:2001PNAS...9810244W, doi:10.1073/pnas.171305098, PMC 56946, PMID 11526236
- Yan, Shi; Wang, Chuan-Chao; Zheng, Hong-Xiang; Wang, Wei; Qin, Zhen-Dong; Wei, Lan-Hai; Wang, Yi; Pan, Xue-Dong; et al. (August 29, 2014). "Y Chromosomes of 40% Chinese Descend from Three Neolithic Super-Grandfathers". PLOS ONE. 9 (8). e105691. arXiv:1310.3897. Bibcode:2014PLoSO...9j5691Y. doi:10.1371/journal.pone.0105691. PMC 4149484. PMID 25170956.
- Y Chromosome Consortium "YCC" (2002). "A Nomenclature System for the Tree of Human Y-Chromosomal Binary Haplogroups". Genome Research. 12 (2): 339–348. doi:10.1101/gr.217602. PMC 155271. PMID 11827954.
- Zerjal, Tatiana; Wells, R. Spencer; Yuldasheva, Nadira; Ruzibakiev, Ruslan; Tyler-Smith, Chris (2002). "A Genetic Landscape Reshaped by Recent Events: Y-Chromosomal Insights into Central Asia". The American Journal of Human Genetics. 71 (3): 466–82. doi:10.1086/342096. PMC 419996. PMID 12145751.
- Zhong H, Shi H, Qi XB, Duan Y, Tan PP, Jin L, SU B, Ma RZ (January 2011). "Extended Y chromosome investigation suggests postglacial migrations of modern humans into East Asia via the northern route". Molecular Biology and Evolution. 28 (1): 717–27. doi:10.1093/molbev/msq247. PMID 20837606.
- Zhou, Ruixia; An, Lizhe; Wang, Xunling; Shao, Wei; Lin, Gonghua; Yu, Weiping; Yi, Lin; Xu, Shijian; et al. (2007). "Testing the hypothesis of an ancient Roman soldier origin of the Liqian people in northwest China: a Y-chromosome perspective". Journal of Human Genetics. 52 (7): 584–91. doi:10.1007/s10038-007-0155-0. PMID 17579807.
Further reading
[edit]- 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. Archived from the original (PDF) on June 24, 2023. Retrieved September 10, 2009.
- 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". The American Society of Human Genetics 57th Annual Meeting October 23–27, 2007; San Diego, California (PDF) (Abstract). p. 273. Archived from the original (PDF) on June 26, 2008.
- 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. Archived (PDF) from the original on October 28, 2008. Retrieved October 30, 2008.
- Wells, Spencer (2002). The Journey of Man: A Genetic Odyssey. Princeton University Press. ISBN 978-0-691-11532-0.
journal articles
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External links
[edit]- DNA Tree
- FTDNA R1a Y-chromosome Haplogroup Project
- R1a1a1 and Subclades Y-DNA Project – Background Family Tree DNA R1a1a1
- TMRCA
- Various
- Danish Demes Regional DNA Project: Y-DNA Haplogroup R1a Archived August 20, 2008, 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
Haplogroup R1a
View on GrokipediaHaplogroup R1a (M420) is a Y-chromosome DNA haplogroup defined by the M420 mutation, tracing patrilineal ancestry and exhibiting high frequencies across Eurasia, particularly among Indo-European-speaking populations in Eastern Europe, Central Asia, and South Asia.[1] Its major subclades diversified around 5,000–6,000 years ago, with ancient DNA confirming expansions from the Pontic-Caspian steppe region during the Bronze Age.[2][3]
The European branch, primarily under Z283, dominates in Slavic groups such as Poles and Russians, where it reaches 50–60% frequency, reflecting migrations linked to the Corded Ware culture.[1][4] In contrast, the Asian branch Z93 prevails in Indo-Iranian populations, correlating with the Sintashta and Andronovo archaeological complexes.[1] These patterns underscore R1a's role as a genetic marker for steppe pastoralist dispersals that facilitated the dissemination of Indo-European languages and technologies, though interpretations remain debated in contexts prioritizing cultural over genetic evidence.[2]
Definition and Genetic Characteristics
Defining Markers and Mutation History
Haplogroup R1a is phylogenetically defined by the single nucleotide polymorphism (SNP) M420, a G-to-A transition on the Y-chromosome that distinguishes it from its sister clade R1b-M343.[1] This marker emerged as a branch from the ancestral haplogroup R1-M173, with the split from R1b estimated at approximately 25,000 years ago (95% confidence interval: 21,300–29,000 years ago), coinciding with the Last Glacial Maximum.[1] Basal R1a*-M420 lineages, lacking downstream mutations, are exceedingly rare in modern populations, occurring sporadically in regions such as Iran, the Caucasus, and parts of Europe, suggesting limited survival of pre-diversification carriers.[5] The mutation history of R1a involves a series of subsequent SNPs that structure its subclades, with M417 marking a major bottleneck and diversification event around 5,800 years ago (95% CI: 4,800–6,800 years ago), after which most contemporary R1a chromosomes descend.[1] This is upstream of Z645, which further bifurcates into European-oriented branches like Z283 (including Z282, peaking in Eastern Europe) and Asian-oriented Z93, reflecting a dual westward and eastward expansion from a likely Near Eastern or Iranian core area of initial radiation.[1] Additional markers, such as M458 within European subclades, arose around 7,900 years ago (SD ±2,600 years), with localized coalescence peaks in Central Europe dating to the early Holocene.[5] These age estimates derive from coalescent analyses of haplotype diversity and SNP phylogeny in large Y-chromosome datasets, emphasizing star-like expansions post-M417 rather than gradual accumulation, consistent with demographic bottlenecks followed by rapid population growth.[1] [5] While basal M420 TMRCA aligns with Paleolithic refugia, the subclade mutations postdate the Neolithic, linking R1a diversification to Bronze Age steppe dynamics rather than earlier migrations.[1]Position in Broader Y-Chromosome Phylogeny
Haplogroup R1a, defined by the single nucleotide polymorphism (SNP) M420, represents a primary subclade of R1-M173 within the human Y-chromosome phylogeny. R1-M173 diverged from its sister clade R1b-M343 approximately 22,000 to 25,000 years ago, based on calibrated mutation rates from whole-genome sequencing of diverse Y-chromosomes.[6] This bifurcation occurred downstream of haplogroup R-M207, the defining marker for haplogroup R, which emerged around 27,000 years ago from its parent P-M45.[1] Haplogroup R constitutes one of the two major descendants of P-M45—the other being Q-M242, predominant in Siberian and Native American populations—positioning R1a as a key Eurasian lineage within non-African Y-chromosome diversity.[7] Upstream, P-M45 (also known as P-F1085) derives from K2b2-P331, a subclade of the broader K-M526 macrohaplogroup that diversified during the Upper Paleolithic era around 40,000 years ago.[8] K-M526 itself branches from the HIJK (IJKMNOPS) supergroup, which traces to F-M89, a foundational haplogroup marking early modern human expansions into Eurasia approximately 45,000 to 50,000 years ago.[7] F-M89 descends from CF-P143, splitting from the CT-M168 lineage that signifies the primary Out-of-Africa migration event circa 60,000 years ago, with CT further rooting in the BT-M91 ancestor shared across non-basal African and Eurasian patrilines.[7] This hierarchical structure, refined through high-coverage sequencing and SNP phylogenies, underscores R1a's intermediate position between ancient African basal clades (A and B) and the explosive diversification of post-glacial Eurasian subclades.[6] The phylogenetic placement of R1a highlights its co-occurrence with R1b as "companion" haplogroups dominating modern European frequencies, though R1a predominates eastward while R1b does westward, reflecting differential migratory expansions rather than a shared basal origin bias.[6] Ongoing refinements to the Y-tree, incorporating thousands of SNPs via projects like the 1000 Genomes and ancient DNA integrations, continue to calibrate branch lengths but affirm R1a's stable topology under R-P231, with no evidence of paraphyly or re-rooting in recent updates.[7]Phylogeny and Subclade Structure
Overall Phylogenetic Topology
Haplogroup R1a (defined by the single-nucleotide polymorphism, or SNP, M420) occupies a basal position within the R1 (M173) branch of the human Y-chromosome phylogeny, with the split from its sister clade R1b (M343) estimated at approximately 25,100 years ago (95% confidence interval: 21,300–29,000 years ago).[1] The TMRCA for R1a-M420 itself is inferred to be older, around 15,000–22,000 years ago based on coalescent modeling of downstream markers, though direct estimates vary due to limited basal lineages.[1] Rare paraphyletic basal R1a* lineages (M420* but negative for downstream markers like SRY10831.2) persist at low frequencies, primarily in Iran and eastern Turkey, suggesting an ancient origin in the Near East or adjacent regions before major post-glacial expansions.[1] The core structure of R1a is dominated by the derived M417 subclade, which encompasses over 99% of modern R1a chromosomes and exhibits a star-like phylogeny indicative of a bottleneck followed by rapid diversification.[1] This clade's TMRCA is approximately 6,800 years ago, with its immediate descendants splitting into two geographically distinct major branches: Z282 (predominant in Europe) and Z93 (predominant in Central and South Asia), both emerging around 5,800 years ago (95% CI: 4,800–6,800 years ago).[1] These bifurcations align with archaeological correlates of Bronze Age expansions, such as the Corded Ware and Andronovo cultures, though phylogenetic resolution alone does not confirm causation. Minor parallel branches under M417, such as L664 (with a TMRCA around 4,700 years ago), occur sporadically in northwestern Europe (e.g., British Isles) and parts of South Asia, representing relictual or secondary dispersals but comprising less than 1% of total R1a diversity.[9] Downstream from Z282, European subclades like Z284 (northwestern Europe), M458 (central-eastern Europe), and Z280/Z283 (eastern Europe and Baltics) show further regional structuring, often with STR haplotype clusters reflecting patrilocal expansions.[1] Similarly, Z93 diversifies into Asian-specific lineages such as Z2124/Z2125 (Central Asia), L657 (South Asia), and M780 (Iranian plateau), with elevated diversity in the Altai region and Indus periphery indicating localized bottlenecks and serial founder effects.[1] Overall, the topology underscores a dual Eurasian expansion model, with limited back-migration evidenced by asymmetric clade distributions and low basal lineage survival outside origin zones; refinements since 2015 via next-generation sequencing have added hundreds of SNPs but preserved this binary framework without altering basal splits.[1][10]Ancestral Haplogroup R and R1 Branches
Haplogroup R (defined by the SNP M207) constitutes a foundational Y-chromosome lineage descending from haplogroup P, with its TMRCA estimated at approximately 30,000 years before present based on aggregated SNP and STR data from modern populations.[8] Ancient DNA evidence, including R* from the ~24,000-year-old Mal'ta-Buret' culture individual in southern Siberia, supports an origin in North Asia during the Upper Paleolithic, potentially linked to Ancient North Eurasian populations that contributed ancestry to both Europeans and Native Americans.[11] This haplogroup's early diversification occurred amid post-Last Glacial Maximum repopulation dynamics, with subsequent migrations facilitating its spread across Eurasia. The primary branches of R are R1 (M173) and R2 (M479). R2 remains concentrated in South Asia, particularly among Dravidian-speaking groups, suggesting limited expansion beyond the Indian subcontinent.[12] In contrast, R1 underwent broader dispersal, with its TMRCA dated to around 20,000 years before present, likely originating in Central or North Asia following a southern refugium phase during the LGM.[13] Y-STR diversity analyses indicate R1's initial post-glacial movements involved northern routes into East Asia around 18,000 years ago, predating major Neolithic expansions.[14] R1 further bifurcated into R1a (M420) and R1b (M343), lineages that dominate modern Y-chromosome variation in much of Eurasia. R1b's core subclades, such as those under M269, exhibit West Asian phylogenetic roots and Holocene founder effects tied to pastoralist expansions, while R1a traces to steppe-associated dispersals.[12] These splits, estimated between 15,000 and 22,000 years ago via mutation rate calibrations, reflect adaptive responses to climatic amelioration and resource availability, enabling R1-derived groups to populate diverse ecological niches from Europe to South Asia.[11] Basal R1* paragroups, though rare today, underscore the lineage's deep antiquity and minimal survival outside major subclades.Core R1a-M420 and Its Immediate Descendants
Haplogroup R1a is defined by the single nucleotide polymorphism (SNP) M420 on the Y-chromosome, marking its divergence from the ancestral haplogroup R1 (defined by M173).[1] Time to most recent common ancestor (TMRCA) estimates for R1a-M420 range from approximately 15,000 to 22,000 years before present, based on SNP accumulation rates and haplotype diversity analyses.[15] Basal paragroup R1a*-M420, lacking derived mutations in downstream branches, occurs at low frequencies and has been identified primarily in samples from Iran and eastern Turkey, suggesting persistence of early lineages in the Near East.[1] The immediate phylogenetic descendants of R1a-M420 include rare basal subclades such as R1a1-SRY10831.2*, which also show limited distribution in the Near East and exhibit low diversity indicative of genetic drift or bottlenecks.[1] However, over 99% of extant R1a chromosomes belong to the major subclade R1a-M417 (also denoted as Page7 or associated with Z645 in refined nomenclature), with a TMRCA estimated at around 5,800 years ago (95% confidence interval: 4,800–6,800 years).[1] [16] This subclade's rapid expansion correlates with Bronze Age population dynamics, as evidenced by elevated star-like haplotype structures in STR data.[1] R1a-M417 represents the core of modern R1a diversity, serving as the progenitor for subsequent radiations, though its own basal lineages remain scarce outside the initial diversification zone near present-day Iran.[1] Phylogenetic resolution beyond M420 relies on high-coverage sequencing, revealing that earlier broad categorizations underestimated the antiquity and regional anchoring of these foundational branches.[1] Ancient DNA from pre-Bronze Age contexts rarely carries R1a-M420, underscoring its relatively recent demographic prominence despite deep origins.[1]Major European Subclades (e.g., Z282)
Haplogroup R1a-Z282 constitutes the primary European branch of R1a-M417, accounting for more than 96% of European R1a-M417 lineages.[1] Its time to most recent common ancestor (TMRCA) is estimated at approximately 5800 years before present (circa 3800 BCE), with a 95% confidence interval of 4800–6800 years ago, based on SNP-based coalescent analysis of 16 Y-STR loci across diverse samples.[1] This subclade exhibits strong geographic localization within Europe, with paragroup R1a-Z282* frequencies reaching about 20% in northern Ukraine, Belarus, and adjacent Russian regions, indicating early diversification centers in Eastern Europe.[1] Ancient DNA evidence links Z282-derived lineages to the Corded Ware culture around 4600 years before present (circa 2600 BCE), supporting associations with Bronze Age expansions in Central and Northern Europe.[1] The major subclades under Z282 include Z284, M458, and Z280, each showing distinct distributions tied to prehistoric population movements. R1a-Z284 is largely confined to Northwest Europe, peaking at approximately 20% in Norway where it comprises the majority of R1a chromosomes (24 out of 26 sampled), and occurs at low frequencies elsewhere, such as single instances in Eastern Europe.[1] This pattern suggests a post-Bronze Age maritime or northern dispersal, potentially linked to Germanic expansions. R1a-M458 predominates in Central-Eastern Europe, with frequencies exceeding 20% in the Czech Republic, Slovakia, Poland, and western Belarus, and ranging from 11% to 33% across broader Slavic regions; it correlates with West Slavic populations and later medieval dispersals.[1] R1a-Z280, encompassing subgroups like M558, displays high diversity and prevalence in Eastern Europe, often exceeding 20% in Baltic and East Slavic areas, with elevated variance pointing to ancient roots.[1] It is frequently associated with Balto-Slavic linguistic groups and Bronze Age cultures extending from the Corded Ware horizon eastward.[1] These subclades collectively underscore Z282's role in Indo-European expansions, with spatial frequency gradients aligning with archaeological evidence of steppe-derived migrations into Europe rather than local Neolithic origins.[1]Major Asian Subclades (e.g., Z93)
Haplogroup R1a-Z93 constitutes the principal Asian branch of R1a-M420, diverging from its European counterpart Z282 around 5,800 years before present through the defining Z93 mutation. This subclade predominates in Central Asia, South Asia, and parts of Southwest Asia, with the paragroup Z93* exhibiting frequencies exceeding 30% among South Siberian Altaians and notable presence (up to 6%) in Kyrgyzstan and Afghanistan. Its expansion correlates with Bronze Age steppe migrations, as evidenced by ancient DNA from Sintashta culture remains dated circa 2100 BCE, which include R1a-Z93 lineages linked to eastward dispersals from the Pontic-Caspian region.[1][17][1] The core diversity of R1a-Z93 resides in its immediate descendant Z94, which encapsulates the bulk of Central and South Asian R1a variation and further ramifies into regionally specific subclades. Prominent among these is L657, a downstream branch under Z94 that achieves high frequencies in the Indian subcontinent, particularly among northern Indo-European-speaking populations, reflecting post-Bronze Age admixture and local expansions estimated around 4,400 years ago. Complementing L657, Z2124—another Z94 subclade—predominates in Iranian and Afghan Pashtun groups, with ancient attestations in Iron Age contexts like Hun-period samples from the 5th century CE, underscoring recurrent nomadic incursions into Southwest and South Asia.[18] Additional Z93 subclades, such as Z2125, contribute to patchy distributions in Central Asian Turkic and Iranian-speaking groups, often tied to Scythian-Saka horizon movements between 900 and 200 BCE. Overall, Z93's phylogenetic structure reveals a star-like expansion pattern post-2500 BCE, with minimal basal Z93* survival outside Siberian isolates, indicating rapid diversification during Andronovo-related spreads across the Eurasian interior. Genetic diversity metrics, including haplotype variance, peak in southern Central Asia, supporting a northerly origin followed by southward gene flow rather than autochthonous South Asian genesis.[18][1]Minor and Peripheral Branches
In addition to the dominant lineages descending from Z645, haplogroup R1a includes several minor and peripheral branches that represent early divergences or rare subclades with restricted distributions. These encompass basal paragroups such as R1a*-M420(xSRY10831.2), identified in 24 males across surveyed populations, with 18 samples from Iran and 3 from eastern Turkey, indicating a concentration in Southwest Asia.[1] Similarly, the upstream R1a1*-SRY10831.2(xM198) clade appears in 6 individuals, comprising 5 Iranians and 1 Kabardin from the Caucasus, underscoring its peripheral occurrence outside major Eurasian expansions.[1] Downstream of M198/M417 but basal to Z282 and Z93, the paragroup Page7*(xZ282,Z93) totals 12 samples in the dataset, reflecting low diversity and frequency suggestive of limited demographic success compared to core branches.[1] Distinct rare markers within these peripheral structures include M204, detected in a single Iranian sample, and M560, observed in 4 individuals: 2 among Burushaski speakers in northern Pakistan, 1 Hazara from Afghanistan, and 1 Iranian Azeri.[1] These lineages exhibit haplotype diversity patterns consistent with ancient persistence in localized refugia rather than large-scale migrations.[1] The geographic clustering of these minor branches, predominantly in Iran and adjacent regions, contrasts sharply with the broad Indo-European-associated spreads of Z282 and Z93, implying they survived as relict populations amid subsequent replacements or dilutions by expanding subclades.[1] TMRCA estimates for basal R1a-M420 exceed 15,000 years, predating the Bronze Age radiations, though precise dating for these peripherals remains tentative due to sparse sampling.[1]Origins and Early Diversification
Primary Origin in the Pontic-Caspian Steppe
Haplogroup R1a-M420, defined by the M420 mutation, is estimated to have originated approximately 15,000–25,000 years ago, with its major subclade M417 exhibiting a time to most recent common ancestor (TMRCA) around 5,500 years before present, aligning with the early Bronze Age in the Pontic-Caspian steppe region.[19] This temporal coincidence supports the steppe as the locus of early diversification, where pastoralist societies transitioned to mobile herding economies conducive to rapid haplogroup expansion. Phylogenetic analyses indicate that basal R1a lineages show patterns of post-glacial coalescent times, with initial radiation likely in Eastern European or adjacent steppe-forest zones rather than distant refugia.[5] Ancient DNA evidence reinforces the Pontic-Caspian steppe as the primary homeland, with R1a-M417 appearing prominently in Corded Ware culture samples (circa 2900–2350 BCE), which derive substantial ancestry from Yamnaya-related steppe populations despite Yamnaya itself predominantly carrying R1b-Z2103.[2] Although absent in core Yamnaya burials, R1a is attested in contemporaneous or slightly later steppe-forest groups like Fatyanovo-Balanovo, suggesting the haplogroup's presence in the broader ecological zone encompassing the western and eastern Pontic-Caspian grasslands.[20] Further east, Srubnaya-Alakulskaya culture individuals (circa 2000–1500 BCE) from the eastern Pontic-Caspian steppe carried R1a, evidencing a major Bronze Age expansion originating there.[21][22] While some phylogenetic reconstructions, such as those by Underhill et al., propose diversification of R1a subclades near present-day Iran based on STR diversity and basal lineages, ancient DNA distributions prioritize the steppe for the explosive growth of both European (Z282) and Asian (Z93) branches, as Iranian samples lack early R1a-M417 equivalents.[23][24] This causal linkage—steppe ecology enabling horse domestication, wheeled vehicles, and hierarchical warrior societies—facilitated R1a's demographic success, distinguishing it from more sedentary Near Eastern populations where equivalent haplogroups did not similarly proliferate.[21]Ancient DNA Evidence Supporting Steppe Homeland
Ancient DNA studies have revealed the presence of haplogroup R1a in Bronze Age populations directly linked to the Pontic-Caspian steppe and adjacent forest-steppe zones, providing empirical support for its origin in this region. In the Corded Ware culture (approximately 2900–2350 BCE), which spanned from the Rhine River to the middle Volga, 10 out of 11 male individuals analyzed from sites in Germany, Poland, and Lithuania carried R1a, predominantly the Z282 subclade.[2] These samples exhibit substantial autosomal ancestry from Yamnaya-related steppe pastoralists (up to 75%), indicating gene flow or cultural continuity from the steppe core, where early R1a diversification likely occurred.[2] To the east, the Sintashta culture (circa 2200–1800 BCE) in the southern Ural steppe yielded R1a-Z93 in nearly all sampled males (9 out of 10), marking the earliest secure instances of this Asian subclade.[25] Sintashta populations show a mixture of Corded Ware-like and local steppe ancestry, with Y-chromosome dominance of R1a suggesting rapid male-mediated expansion within the steppe environment, contemporaneous with innovations in spoke-wheeled chariots and fortified settlements. This pattern aligns with phylogenetic estimates placing the split between European (Z282) and Asian (Z93) R1a branches around 3500–3000 BCE in the steppe vicinity.[26] Later Bronze Age steppe groups, such as the Srubnaya-Alakul horizon (circa 1900–1200 BCE) in the eastern Pontic-Caspian region, further document R1a prevalence, with multiple individuals carrying the haplogroup amid evidence of demographic expansion.[21] Genome-wide data from these samples indicate a stable genetic profile tracing back to earlier steppe sources, predating dispersals into Europe and Asia.[21] The temporal and geographic clustering of basal R1a-M420 derivatives in these contexts, absent in contemporaneous non-steppe Neolithic farmers or hunter-gatherers, underscores the steppe as the primary homeland, with coalescent times for major subclades (circa 5000–6000 years ago) matching archaeological timelines of pastoralist mobility.[2][25]Evaluation of Alternative Origin Hypotheses
One prominent alternative hypothesis posits the Indian subcontinent as the primary origin of haplogroup R1a, particularly its M417 subclade, based on elevated microsatellite (STR) diversity and frequency among certain Indian populations, such as Brahmins, suggesting an autochthonous development predating Indo-European migrations.[27][28] Proponents argued that this diversity indicated a local expansion over millennia, challenging Steppe-derived introductions and aligning with cultural narratives of indigenous Aryan origins.[27] However, this view has been undermined by SNP-based phylogenetic reconstructions, which estimate the time to most recent common ancestor (TMRCA) for R1a-M417 at approximately 5,800 years before present, with major branches Z282 (predominantly European) and Z93 (predominantly Asian) diverging around 4,700–5,200 years ago—timings incompatible with a deep South Asian root and instead aligning with Bronze Age Steppe cultures.[23] STR variance, once central to Indian origin claims, proves unreliable for pinpointing origins due to its sensitivity to population size and serial founder effects rather than true antiquity, as later high-resolution SNP and whole-genome studies demonstrate younger coalescent times for Indian R1a-Z93 lineages compared to European counterparts.[23][29] Ancient DNA evidence further refutes a pre-Bronze Age Indian origin, as no R1a lineages appear in Indus Valley Civilization samples (circa 2600–1900 BCE), which instead carry haplogroups like H and L without Steppe ancestry components.[29] In contrast, R1a-M417 derivatives, including Z93, emerge in Steppe-associated contexts like Sintashta (circa 2100–1800 BCE), predating their detection in South Asia, where Z93 frequencies correlate with male-biased Steppe ancestry influx around 2000–1500 BCE, disproportionately elevated in upper-caste groups.[29] This pattern indicates unidirectional gene flow from the Steppe, not vice versa. Other proposed alternatives, such as an Iranian Plateau or Anatolian origin, similarly lack substantiation; R1a frequencies remain low in ancient Near Eastern samples, and basal diversity does not cluster there, with phylogenetic gradients pointing eastward to the Pontic-Caspian region rather than southward or westward.[23] These hypotheses often stem from outdated diversity metrics or selective modern sampling, overlooking causal dynamics like bottleneck-driven expansions from small Steppe founder populations, which better explain the haplogroup's star-like phylogeny and rapid dispersal. Empirical genomic data, prioritizing ancient sequences over interpretive modern variances, thus overwhelmingly favor the Steppe as the locus of R1a-M420's defining diversification, rendering alternatives inconsistent with the chronological and geographic evidence.[23][29]Prehistoric Migrations and Expansions
Initial Bronze Age Dispersals (Yamnaya and Corded Ware)
The Yamnaya culture, dating from circa 3300 to 2600 BCE in the Pontic-Caspian steppe, primarily carried Y-haplogroup R1b-M269, with subclade Z2103 dominating male lineages in ancient DNA samples from over 100 individuals, while R1a remains rare or undetected in core Yamnaya burials.[30] This R1b dominance reflects a patrilineal continuity from preceding Eastern Hunter-Gatherer populations, but the culture's expansive pastoralist lifestyle facilitated the initial dissemination of steppe autosomal ancestry westward into Europe and eastward into Central Asia, laying genetic foundations for subsequent Bronze Age groups without substantial R1a contribution from Yamnaya itself. Although direct R1a dispersal via Yamnaya is minimal, the culture's migrations correlate with the broader Bronze Age steppe expansions that enabled related populations carrying R1a to emerge and spread, as seen in peripheral or successor groups like those in the Don-Volga region where pre-Corded Ware R1a instances appear sporadically before 2900 BCE.[31] Ancient DNA evidence indicates that R1a-M417 and its derivatives, such as Z283, were present in Eastern European contexts prior to major Yamnaya movements but gained prominence in the formation of derivative cultures rather than the Yamnaya core.[2] The Corded Ware culture, emerging around 2900 BCE and extending to circa 2350 BCE across Central, Northern, and Eastern Europe, represents the primary vector for R1a dispersal during the early Bronze Age, with ancient DNA from dozens of male samples—particularly from sites in Poland, Germany, and the Baltic region—revealing R1a frequencies exceeding 70% in early phases, often subclade R1a-Z283.[32] [33] This culture incorporated substantial Yamnaya-derived steppe ancestry (up to 75% in some models) through male-mediated migration, effectively replacing Neolithic farmer Y-lineages like G2a and I2 in northern Europe, while R1a bearers expanded from probable origins near the middle Dnieper or upper Volga, driving linguistic and cultural shifts associated with Indo-European branches. [2] Genetic continuity links Corded Ware R1a to later Baltic and Slavic populations, underscoring its role in the initial Bronze Age peopling of temperate Europe.[32]Sintashta and Andronovo Culture Associations
The Sintashta culture, dated to approximately 2200–1800 BCE and centered in the southern Ural steppe region extending into northern Kazakhstan, is distinguished by its fortified hilltop settlements, advanced bronze metallurgy, and the earliest archaeologically attested spoke-wheeled chariots, which facilitated enhanced mobility for pastoralist warriors. Ancient DNA from 101 Bronze Age Eurasian samples, including those from Sintashta sites, revealed that male individuals from this culture carried Y-haplogroup R1a, with subclade Z93 (specifically under R1a1a1b2-Z93) identified in analyzed burials such as RISE392. This genetic signature reflects a synthesis of incoming western steppe ancestry—linked to earlier Corded Ware populations—with minor local admixtures, marking Sintashta as a pivotal eastern extension of R1a-bearing groups. Sintashta's successor, the Andronovo culture (circa 2000–900 BCE), encompassed a broader expanse across the Eurasian steppes from the Ural Mountains to the Altai region and into Central Asia, characterized by semi-nomadic pastoralism, kurgan burials, and continuity in ceramic and metallurgical traditions. Genetic studies of Andronovo-associated remains consistently show male Y-chromosomes dominated by R1a-Z93, with frequencies approaching 100% in some Middle to Late Bronze Age steppe samples proxying Andronovo horizons, as evidenced in datasets from the Volga-Ural and Kazakhstan regions.[34][25] This uniformity in paternal lineages supports models of male-biased expansions, where R1a-Z93 carriers maintained genetic continuity amid geographic dispersal.[34] The prevalence of R1a-Z93 in Sintashta and Andronovo underscores their role in the initial Bronze Age radiations of this subclade eastward from the Pontic-Caspian core, distinct from the Z282 branch predominant in contemporaneous western European expansions.[25] These cultures' genetic profiles align with archaeological evidence of technological and cultural innovations that enabled further migrations into South and Central Asia, though debates persist on the extent of elite dominance versus broader population replacements in recipient regions.[34]Later Iron Age Movements into Asia and Europe
Ancient DNA evidence indicates that during the later Iron Age (ca. 900 BCE–400 CE), populations carrying R1a subclades, particularly under Z93, expanded eastward with Scythian nomads from the Pontic-Caspian steppe into Central Asia and the Altai region, as seen in kurgan burials from Kazakhstan and southern Siberia dated 900–200 BCE, where R1a-Z93 comprised a significant portion of male lineages alongside Q and R1b variants.[35] These movements facilitated the spread of Iranian-speaking steppe pastoralists, with genetic continuity from earlier Andronovo-related groups but increased East Eurasian admixture in eastern samples, suggesting gene flow via marriage alliances rather than wholesale population replacement.[36] Frequencies of R1a-Z93 in these eastern Scythian assemblages reached 40–60% in patrilineal clans, underscoring its role in male-biased dispersals across Asia.[35] In the western steppe, Scythians of the Middle Don region (7th century BCE–1st century CE) exhibited strong patrilineal clustering under R1a-Y2631 (a Z93 descendant), with at least 27 males from a single clan sharing this lineage, indicating clan-based expansions and limited diversity before Sarmatian displacements around the 3rd century BCE.[35] Sarmatians, succeeding Scythians, maintained R1a-Z93/Z94 prominence while incorporating local elements, enabling further eastward pushes into the Volga-Ural zone and southward toward the Caucasus.[23] Westward into Europe, Sarmatian groups penetrated the Carpathian Basin and Danube frontiers by the 1st–5th centuries CE, introducing R1a-Z94 alongside G and J2a haplogroups, as evidenced by genomic analyses of 156 individuals from Hungarian sites showing steppe-derived ancestry and Y-chromosome diversity reflective of nomadic incursions amid Roman provincial interactions.[37] These movements contributed minor R1a influxes to Central European populations, contrasting with predominant local R1b and I2 lineages in pre-Sarmatian Iron Age contexts, and lacked the scale of earlier Bronze Age dispersals.[4] Overall, Iron Age R1a dynamics emphasized Asian-directed expansions via mobile steppe elites, with European gene flow remaining peripheral until later periods.[35]Modern Geographic Distribution
Prevalence in Europe
Haplogroup R1a exhibits its highest frequencies in Europe within Eastern European populations, particularly those associated with Slavic ethnolinguistic groups, where it often constitutes 40-60% of male lineages. In Poland, R1a reaches peaks exceeding 50% nationally, with subclades like M458 contributing significantly to this prevalence.[1] Similarly, in Russia, approximately every second Y-chromosome belongs to R1a, yielding frequencies of 45-50% or higher in central and northern regions.[38] Ukraine and Belarus show comparable levels, with R1a at around 44% and over 50%, respectively, driven by Z282 and M558 subclades.[39][1] Frequencies decline westward and northward from this core. In Central Europe, Czechia and Slovakia register 30-40%, while Hungary averages 20-40%, reflecting partial Slavic admixture.[39] Germany exhibits 15-25%, concentrated in eastern areas, whereas France and the United Kingdom maintain low incidences under 10% and around 5%, respectively, indicating limited Bronze Age Steppe influx in Western Europe.[1] In Scandinavia, Norway stands out with up to 25% due to the Z284 subclade, linked to Viking-era movements, compared to 10-15% in Sweden.[1] Southern Europe generally shows minimal R1a (<10%), except in Albania and Greece where isolated pockets reach 10-20%.[1] These patterns align with ancient DNA evidence of Corded Ware and later Slavic expansions, with R1a-Z283 dominating European subclades (over 90% of continental R1a), contrasting with Asian Z93 branches.[1] Variation within countries underscores regional heterogeneity, such as higher eastern concentrations in Poland and Ukraine.[39]| Country/Region | R1a Frequency (%) | Primary Subclades Noted |
|---|---|---|
| Poland | 50-60 | M458, Z282 |
| Russia | 45-50+ | Z282, M558 |
| Ukraine | ~44 | Z282, M558 |
| Belarus | >50 | M458, M558 |
| Norway | ~25 | Z284 |
| Germany | 15-25 | Z282 |
