The R1a (R-M420) subclade diverged from R1 (R-M173) 15-25,000[2][4][5] years ago, itssubclade 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 ofProto-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 newparagroup (designated R-M420*) for the relatively rare lineages which are not in the R-SRY10831.2 (R1a1) branch leading to R-M17.
Thegenetic divergence of R1a (M420) is estimated to have occurred 25,000[2] years ago, which is the time of thelast 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 theMesolithic inEastern Hunter-Gatherers (from Eastern Europe, c. 13,000 years ago),[7][8] and the earliest case of R* amongUpper PaleolithicAncient North Eurasians,[9] from which the Eastern Hunter-Gatherers predominantly derive their ancestry.[10]
Diversification of R1a1a1 (M417) and ancient migrations
R1a origins (Underhill 2009;[3] R1a1a origins (Pamjav et al. 2012); possible migration R1a to Baltic coast; and R1a1a oldest expansion and highest frequency (Underhill et al. 2014)
Semino et al. (2000) proposedUkrainian origins, and a postglacial spread of the R1a1 haplogroup during theLate Glacial Maximum, subsequently magnified by the expansion of the Kurgan culture into Europe and eastward.[14] Spencer Wells proposes Central Asian origins, suggesting that the distribution and age of R1a1 points to an ancient migration corresponding to the spread by theKurgan people in their expansion from theEurasian steppe.[15] According toPamjav 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.[16]
Three genetic studies in 2015 gave support to theKurgan theory of Gimbutas regarding theIndo-European Urheimat. According to those studies, haplogroupsR1b 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.[17][18][19]
Silva et al. (2017) noted that R1a in South Asia most "likely spread from a singleCentral Asian source pool, there do seem to be at least three and probably more R1a founder clades within theIndian subcontinent, consistent with multiple waves of arrival."[20] According to Martin P. Richards, co-author ofSilva 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."[21][22]
European middle-Neolithic period. Comb Ware culture c. 4200 – c. 2000 BCECorded Ware culture (c. 2900 – c. 2350 BCE
David Anthony considers theYamnaya culture to be theIndo-European Urheimat.[23][24] According toHaak 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 theCorded Ware culture, noting that R1a and R1b may have "spread into Europe from theEast after 3,000 BCE".[25] Yet, all their seven Yamnaya samples belonged to theR1b-M269 subclade,[25] 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.[26]
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.[27]
According toLeo Klejn, the absence of haplogroup R1a in Yamnaya remains (despite its presence in Eneolithic Samara andEastern Hunter Gatherer populations) makes it unlikely that Europeans inherited haplogroup R1a from Yamnaya.[28]
ArchaeologistBarry 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.[29]
Kivisild et al. (2003) have proposed either South orWest Asia,[31][note 3] whileMirabal et al. (2009) see support for both South and Central Asia.[12] Sengupta et al. (2006) have proposed Indian origins.[32] Thanseem et al. (2006) have proposed either South or Central Asia.[33] Sahoo et al. (2006) have proposed either South or West Asia.[34] Thangaraj et al. (2010) have also proposed a South Asian origin.[35] 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 highestSTR diversity within R1a1a,[36][37][12][3][1][38] and subsequent olderTMRCA 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 theSaharia tribe, a scheduled caste of theBundelkhand region ofCentral India.[1][38]
From these findings some researchers argued that R1a1a originated in South Asia,[37][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.[37][36][3][note 6]
The conclusion that R1a originated in India has been questioned by more recent research,[20][40][note 7] offering proof that R1a arrived in India with multiple waves of migration.[20][41]
Proposed Transcaucasia and West Asian origins and possible influence on Indus Valley Civilization
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 theBalkans.[note 8] TheRössen culture (4,600–4,300 BC), which was situated onGermany 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 toIndia viaIran[43] and expanded there during theIndus Valley civilization.[2][44]
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 fromTranscaucasia intoSouth Asia",[43] noting that such an expansion is compatible with "the archeological records of eastward expansion ofWest Asian populations in the 4th millennium BCE culminating in the so-calledKura-Araxes migrations in the post-Uruk IV period."[43] Yet, Lazaridis noted that sample I1635 ofLazaridis et al. (2016), theirArmenian Kura-Araxes sample, carried Y-haplogroup R1b1-M415(xM269)[note 10] (also called R1b1a1b-CTS3187).[45][unreliable source?]
According toUnderhill 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."[44][note 13]
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).
The topology of R1a is as follows (codes [in brackets] non-isogg codes):[11][48][verification needed][49][2][50] Tatiana et al. (2014) "rapid diversification process ofK-M526 likely occurred inSoutheast Asia, with subsequent westward expansions of the ancestors of haplogroupsR andQ."[51]
R1a is distinguished by several unique markers, including the M420 mutation. It is a subclade ofHaplogroup R-M173 (previously called R1). R1a has the sister-subcladesHaplogroup R1b-M343, and the paragroup R-M173*.
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 theVasylivka, Bakhmut Raion, Donetsk Oblast.[54][5]
R1a2 (R-YP4141) has two branches R1a2a (R-YP5018) and R1a2b (R-YP4132).[55]
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][56]
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 1/15Himachal Pradesh Rajput samples.[37] Underhill et al. (2009) reported 1/51 inNorway, 3/305 inSweden, 1/57Greek Macedonians, 1/150 (or 2/150) Iranians, 2/734 ethnicArmenians, 1/141Kabardians, 1/121Omanis, 1/164 in theUnited Arab Emirates, and 3/612 inTurkey. 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-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 theCoptic community in Egypt respectively.[57]
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-M458 is a mainlySlavic SNP, characterized by its own mutation, and was first calledcluster N. Underhill et al. (2009) found it to be present in modern European populations roughly between theRhine catchment and theUral 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 inUsedom, Mecklenburg-Vorpommern, Germany.[59] The paper by Underhill et al. (2009) also reports a surprisingly high frequency of M458 in someNorthern Caucasian populations (18% amongAk Nogai,[60] 7.8% amongQara Nogai and 3.4% amongAbazas).[61]
R1a1a1b1a1a (R-L260), commonly referred to asWest Slavic orPolish, is a subclade of the larger parent group R-M458, and was first identified as an STR cluster byPawlowski et al. 2002. In 2010 it was verified to be a haplogroup identified by its own mutation (SNP).[62] It apparently accounts for about 8% of Polish men, making it the most common subclade in Poland. Outside of Poland it is less common.[63] In addition to Poland, it is mainly found in theCzech Republic andSlovakia, 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 theIron Age, with significant population expansion less than 1,500 years ago.[64]
R-M334 ([R1a1a1g1],[50] a subclade of [R1a1a1g] (M458)[50] c.q. R1a1a1b1a1 (M458)[49]) was found by Underhill et al. (2009) only in one Estonian man and may define a very recently founded and small clade.[3]
R1a1a1b1a2b3* (M417+, Z645+, Z283+, Z282+, Z280+, CTS1211+, CTS3402, Y33+, CTS3318+, Y2613+) (Gwozdz's Cluster K)[48][verification needed] is a STR based group that is R-M17(xM458). This cluster is common in Poland but not exclusive to Poland.[64]
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, inNaumovskoye, andKhanevo, 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-M434 (R1a1a6) is a subclade of Z2125. It was detected in 14 people (out of 3667 people tested), all in a restricted geographical range fromPakistan toOman. This likely reflects a recent mutation event in Pakistan.[3]
R-M560 is very rare and was only observed in four samples: two Burushaski speakers (north Pakistan), one Hazara (Afghanistan), and one Iranian Azerbaijani.[2]
R-M780 (R1a1b2a2*) occurs at high frequency in South Asia: India, Pakistan, Afghanistan, and the Himalayas. Turkey share R1a (12.1%) sublineages.[65] Roma from Slovakia share 3% of R1a[66] The group also occurs at >3% in some Iranian populations and is present at >30% in Roma from Croatia and Hungary.[2]
In Mesolithic Europe, R1a is characteristic ofEastern Hunter-Gatherers (EHGs).[67] A male EHG of theVeretye culture buried atPeschanitsa nearLake Lacha inArkhangelsk Oblast, Russia c. 10,700 BCE was found to be a carrier of the paternal haplogroup R1a5-YP1301 and the maternal haplogroupU4a.[68][69][67] A male, named PES001, fromPeschanitsa innorthwestern 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 fromMinino in northwestern Russia.[70] A Mesolithic male fromKarelia c. 8,800 BCE to 7950 BCE has been found to be carrying haplogroup R1a.[71] AMesolithic male buried atDeriivka c. 7000 BCE to 6700 BCE carried the paternal haplogroup R1a and the maternalU5a2a.[19] Another male from Karelia from c. 5,500 to 5,000 BC, who was considered an EHG, carried haplogroup R1a.[17] A male from theComb Ceramic culture inKudruküla c. 5,900 BCE to 3,800 BCE has been determined to be a carrier of R1a and the maternalU2e1.[72] According to archaeologist David Anthony, the paternalR1a-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)."[73] R1a has been found in theCorded Ware culture,[74][75] in which it is predominant.[76] Examined males of the Bronze AgeFatyanovo culture belong entirely to R1a, specifically subclade R1a-Z93.[67][68][77]
Haplogroup R1a has later been found in ancient fossils associated with theUrnfield culture;[78] as well as the burial of the remains of theSintashta,[18]Andronovo,[79] thePazyryk,[80]Tagar,[79]Tashtyk,[79] andSrubnaya cultures, the inhabitants of ancientTanais,[81] in theTarim mummies,[82] and the aristocracy ofXiongnu.[83] The skeletal remains of a father and his two sons, from an archaeological site discovered in 2005 near Eulau (inSaxony-Anhalt,Germany) and dated to about 2600 BCE, tested positive for the Y-SNP marker SRY10831.2. TheYsearch 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 widespreadCorded Ware culture.[74]
In Europe, the R1a1a sub-clade is primarily characteristic of Balto-Slavic populations, with two exceptions: southern Slavs and northern Russians.[84] The highest frequency of R1a1a in Europe is observed inSorbs (63%),[85] aWest Slavic ethnic group, followed byHungarians (60%).[14] Other groups with significant R1a1a, ranging from 27% to up to 58%, includeCzechs,Poles,Slovenians,Slovaks,Moldovans,Belarusians,Rusyns,Ukrainians, andRussians.[84][85][14] 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 theBaltics, R1a1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).[86][87][88][14][89]
There is also a significant presence in peoples ofGermanic descent, with highest levels inNorway,Sweden andIceland, where between 20 and 30% of men are in R1a1a.[90][91]Vikings andNormans may have also carried the R1a1a lineage further out, accounting for at least part of the small presence in theBritish Isles, theCanary Islands, andSicily.[92][93] Haplogroup R1a1a averages between 10 and 30% inGermans, with a peak inRostock at 31.3%.[94] R1a1a is found at a very low frequency amongDutch people (3.7%)[14] and is virtually absent inDanes.[95]
R1a is virtually composed only of the Z284 subclade inScandinavia. 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 inTurkey, 12,1%[65][2]West Slavs andHungarians 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] TheBalts,East Slavs,Serbs,Macedonians,Bulgarians andRomanians 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 detailedphylogeny of the subclades.[103][104] 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 theEurasian steppe according to archaeological and toponymic references.[note 15]
Haber et al. (2012) found R1a1a-M17 in 26.0% (53/204) of a set of samples fromAfghanistan, including 60% (3/5) of a sample ofNuristanis, 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.[106]
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 fromBaghlan and 29/53 or 54.7% of a sample of Pashtuns fromKunduz), 29.1% (37/127) of a pool of samples of Uzbeks from Afghanistan (including 28/94 or 29.8% of a sample of Uzbeks fromJawzjan, 8/28 or 28.6% of a sample of Uzbeks fromSar-e Pol, and 1/5 or 20% of a sample of Uzbeks fromBalkh), 27.5% (39/142) of a pool of samples of Tajiks from Afghanistan (including 22/54 or 40.7% of a sample of Tajiks fromBalkh, 9/35 or 25.7% of a sample of Tajiks fromTakhar, 4/16 or 25.0% of a sample of Tajiks fromSamangan, and 4/37 or 10.8% of a sample of Tajiks fromBadakhshan), 16.2% (12/74) of a sample ofTurkmens fromJawzjan, and 9.1% (7/77) of a pair of samples ofHazara from Afghanistan (including 7/69 or 10.1% of a sample of Hazara fromBamiyan and 0/8 or 0% of a sample of Hazara fromBalkh).[107]
Ashirbekov et al. (2017) found R1a-M198 in 6.03% (78/1294) of a set of samples ofKazakhs fromKazakhstan. 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.[109]
Besides these, studies show high percentages in regionally diverse groups such asManipuris (50%)[3] to the extreme North East and amongPunjabis (47%)[31] to the extreme North West.
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 ofDolan 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.[117] Note that Darya Boyi Village is located in a remote oasis formed by theKeriya River in theTaklamakan Desert. A 2011 Y-DNA study found Y-dna R1a1 in 10% of a sample of southernHui people from Yunnan, 1.6% of a sample ofTibetan people fromTibet (Tibet Autonomous Region), 1.6% of a sample ofXibe people from Xinjiang, 3.2% of a sample of northern Hui fromNingxia, 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 ofMongols fromInner Mongolia. A different subclade of R1 was also found in 1.5% of a sample of northern Hui from Ningxia.[118] in the same study there were no cases of R1a detected at all in 6 samples of Han Chinese inYunnan, 1 sample of Han inGuangxi, 5 samples of Han inGuizhou, 2 samples of Han inGuangdong, 2 samples of Han inFujian, 2 samples of Han inZhejiang, 1 sample of Han inShanghai, 1 samples of Han inJiangxi, 2 samples of Han inHunan, 1 sample of Han inHubei, 2 samples of Han inSichuan, 1 sample of Han inChongqing, 3 samples of Han inShandong, 5 samples of Han inGansu, 3 samples of Han inJilin and 2 samples of Han inHeilongjiang.[119] 40% of Salars, 45.2% ofTajiks of Xinjiang, 54.3% of Dongxiang, 60.6% ofTatars and 68.9% ofKyrgyz 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.[120]
Y-haplogroups R1a-M420 and R2-M479 are found inEde (8.3% and 4.2%) andGiarai (3.7% and 3.7%) peoples inVietnam. TheCham additionally have haplogroups R-M17 (13.6%) andR-M124 (3.4%).
R1a1a1b2a2a (R-Z2123) and R1a1 are found inKhmer peoples fromThailand (3.4%) andCambodia (7.2%) respectively. Haplogroup R1a1a1b2a1b (R-Y6) is also found amongKuy 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.[122]
R1a1a has been found in various forms, in most parts ofWestern Asia, in widely varying concentrations, from almost no presence in areas such asJordan, to much higher levels in parts ofKuwait andIran. The Shimar (Shammar) Bedouin tribe inKuwait show the highest frequency in the Middle East at 43%.[123][124][125]
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 ofTehran andIsfahan.Regueiro 2006 in a study ofIran, noted much higher frequencies in the south than the north.
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 fromTehran 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).[107]
Malyarchuk et al. (2013) found R1a1-SRY10831.2 in 20.8% (16/77) of a sample ofPersians collected in the provinces ofKhorasan andKerman in eastern Iran, but they did not find any member of this haplogroup in a sample of 25Kurds collected in the province ofKermanshah in western Iran.[108]
Further to the north of these Western Asian regions on the other hand, R1a1a levels start to increase in theCaucasus, 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 theKarachay-Balkar language among whom about one quarter of men tested so far are in haplogroup R1a1a.[3]
The historic naming system commonly used for R1a was inconsistent in different published sources, because it changed often; this requires some explanation.
In 2002, theY 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 ofSNP mutations can also be used to name clades or haplogroups. For example, as M173 is currently thedefining 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 andISOGG 2012) R1a is now defined by the M420 mutation: in this updated tree, the subclade defined by SRY1532.2 has moved from R1a to R1a1, and Eu19 (R-M17) from R1a1 to R1a1a.
More recent updates recorded at the ISOGG reference webpage involve branches of R-M17, including one major branch, R-M417.
Contrasting family trees for R1a, showing the evolution of understanding of this clade
^Van Oven M, Van Geystelen A, Kayser M, Decorte R, Larmuseau HD (2014). "Seeing the wood for the trees: a minimal reference phylogeny for the human Y chromosome".Human Mutation.35 (2):187–91.doi:10.1002/humu.22468.PMID24166809.S2CID23291764.
^K-M2313*, which as yet has no phylogenetic name, has been documented in two living individuals, who have ethnic ties to India and South East Asia. In addition, K-Y28299, which appears to be a primary branch of K-M2313, has been found in three living individuals from India. See: Poznikop. cit.;YFull YTree v5.08, 2017, "K-M2335", and;PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (Access date of these pages: 9 December 2017)
^ Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)
^ Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)
Mathieson, Iain (2015). "Eight thousand years of natural selection in Europe".bioRxiv10.1101/016477.
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.
^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."[31]
^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.[39] *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."
^ThoughSengupta (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."[36]
^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."[40]
^Yet, Haak et al. also explicitly state: "a type of Near Eastern ancestry different from that which was introduced by early farmers".[clarification needed][42]
^According to Family Tree DNA, L664 formed 4,700 ybp, that is, 2,700 BCE.[11]
^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)[46]
^According to Family Tree DNA, M780 formed 4700 ybp.[11] This dating coincides with the eastward movement between 2800 and 2600 BCE of the Yamnaya culture into the region of thePoltavka culture, a predecessor of theSintashta culture, from which the Indo-Iranians originated. M780 is concentrated in the Ganges Valley, the locus of the classicVedic 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 theSteppe 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)."[47]
^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.
^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.PMC6822619.PMID31488661.Y chromosome haplogroup types R1b or R1a not represented in Iran and Turan in this period ...
^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.ISBN978-0-3064-6295-5.
^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.ISSN0065-101X.Archived from the original on December 25, 2022. RetrievedNovember 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)."
^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.PMC1380230.PMID16400607."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."
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_and_R1a1a_oldest_expansion_and_highest_frequency_(2014).jpg)
