Haplogroup R-M269#R1b1a1a2a (R-L23)
{{short description|Gene group}}
{{Infobox haplogroup
|name = R-M269
|map = File:Geographical distribution of haplogroup frequency of hgR1b1b2.png
|caption = Projected spatial frequency distribution for haplogroup R-M269 in Europe.Balaresque et al. (2010), figure 1B: "Geographical distribution of haplogroup frequency of hgR1b1b2, shown as an interpolated spatial frequency surface. Filled circles indicate populations for which microsatellite data and TMRCA estimates are available. Unfilled circles indicate populations included to illustrate R1b1b2 frequency only. Population codes are defined in [http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.1000285#pbio-1000285-t001 Table 1]."
|origin-date = 4,000–10,000 BP"Mean estimates for individual populations vary (Table 2), but the oldest value is in Central Turkey (7,989 y [95% confidence interval (CI): 5,661–11,014]), and the youngest in Cornwall (5,460 y [3,764–7,777]). The mean estimate for the entire dataset is 6,512 y (95% CI: 4,577–9,063 years), with a growth rate of 1.95% (1.02%–3.30%). Thus, we see clear evidence of rapid expansion, which cannot have begun before the Neolithic period." Balaresque et al. (2010).
|origin-place = controversial;{{cite journal |last1=Villaescusa |first1= Patricia| title = Characterization of the Iberian Y chromosome haplogroup R-DF27 in Northern Spain | journal = Forensic Science International: Genetics | volume = 27 | issue = | pages = 142–148| date =2017 | doi = 10.1016/j.fsigen.2016.12.013 |pmid= 28073088| quote= The origin of R-M269 has been the subject of great controversy[6], [7], [8], as it was originally believed to have originated in the Palaeolithic [9], [10]. More recent analysis [11], [12] suggested that this lineage had a Neolithic origin, but this claim was challenged [7] due to the Y-STR choice for computing the coalescence times and sample ascertainment. The last studies involving next-generation sequencing (NGS) of the Y-chromosome [8], [13] and the analysis of ancient DNA [14] bring light to the debate, as they support more recent origin and continent-wide expansion of the main European patrilineages, including R-M269 (≈5 KYA, middle Neolithic).| issn =1872-4973 }} most likely North Caucasus, associated with Indo-European migrations{{cite journal | vauthors = Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, Llamas B, Brandt G, Nordenfelt S, Harney E, Stewardson K, Fu Q, Mittnik A, Bánffy E, Economou C, Francken M, Friederich S, Pena RG, Hallgren F, Khartanovich V, Khokhlov A, Kunst M, Kuznetsov P, Meller H, Mochalov O, Moiseyev V, Nicklisch N, Pichler SL, Risch R, Rojo Guerra MA, Roth C, Szécsényi-Nagy A, Wahl J, Meyer M, Krause J, Brown D, Anthony D, Cooper A, Alt KW, Reich D | display-authors = 6 | title = Massive migration from the steppe was a source for Indo-European languages in Europe | journal = Nature | volume = 522 | issue = 7555 | pages = 207–11 | date = June 2015 | pmid = 25731166 | pmc = 5048219 | doi = 10.1038/nature14317 | arxiv = 1502.02783 | bibcode = 2015Natur.522..207H }}{{cite journal |journal=Science |volume=377 |issue=6609 |date=2022 |doi=10.1126/science.abm4247 |title=The genetic history of the Southern Arc: a bridge between West Asia and Europe |last1=Lazaridis |first1=Iosif |display-authors=etal |pages=Supplementary material, p.332 |pmid=36007055 |pmc=10064553 |quote=Given that within the phylogeny of R-M269 (R-PF7562, (R-L51, R-Z2103 is meant) both R-PF7562 and R-Z2103 have their earliest examples in the North Caucasus and steppe to the north, the most likely hypothesis is that the entire R-M269 clade originated there as well, with R-L51 representing a lineage that eventually became highly successful in mainland Europe, R-PF7562 a lineage that did not achieve the prominence of its relatives, and R-Z2103 became highly successful (briefly) as part of the Yamnaya culture and its offshoots}}
|ancestor = R1b1a1a (R-P297)
|descendants = L23; L51/M412, L151/P310; Z2103
|mutations = M269
|members =
}}
Haplogroup R-M269 is the sub-clade of human Y-chromosome haplogroup R1b that is defined by the SNP marker M269. According to ISOGG 2020 it is phylogenetically classified as R1b1a1b. It underwent intensive research and was previously classified as R1b1a2 (2003 to 2005), R1b1c (2005 to 2008), R1b1b2 (2008 to 2011) and R1b1a1a2 (2011 to 2020).dates according to the ISOGG trees for each respective year.
The oldest R-M269 samples have been found in the northern Caucasus region.{{Cite journal |last1=Ghalichi |first1=Ayshin |last2=Reinhold |first2=Sabine |last3=Rohrlach |first3=Adam B. |last4=Kalmykov |first4=Alexey A. |last5=Childebayeva |first5=Ainash |last6=Yu |first6=He |last7=Aron |first7=Franziska |last8=Semerau |first8=Lena |last9=Bastert-Lamprichs |first9=Katrin |last10=Belinskiy |first10=Andrey B. |last11=Berezina |first11=Natalia Y. |last12=Berezin |first12=Yakov B. |last13=Broomandkhoshbacht |first13=Nasreen |last14=Buzhilova |first14=Alexandra P. |last15=Erlikh |first15=Vladimir R. |date=November 2024 |title=The rise and transformation of Bronze Age pastoralists in the Caucasus |journal=Nature |volume=635 |issue=8040 |pages=917–925 |doi=10.1038/s41586-024-08113-5 |issn=1476-4687 |pmid=39478221|pmc=11602729 |bibcode=2024Natur.635..917G }}
R-M269 is of particular interest for the genetic history of Western Europe, being the most common European haplogroup. It increases in frequency on an east to west gradient (its prevalence in Poland estimated at 22.7%, compared to Wales at 92.3%). It is carried by approximately 110 million European men (2010 estimate).{{cite journal | vauthors = Balaresque P, Bowden GR, Adams SM, Leung HY, King TE, Rosser ZH, Goodwin J, Moisan JP, Richard C, Millward A, Demaine AG, Barbujani G, Previderè C, Wilson IJ, Tyler-Smith C, Jobling MA | display-authors = 6 | title = A predominantly neolithic origin for European paternal lineages | journal = PLOS Biology | volume = 8 | issue = 1 | pages = e1000285 | date = January 2010 | pmid = 20087410 | pmc = 2799514 | doi = 10.1371/journal.pbio.1000285 | veditors = Penny D |doi-access=free}}
The age of the mutation M269 is estimated at 4,000 to 10,000 years ago.
Origin
R-M269 had formerly been dated to the Upper Paleolithic,{{cite journal | vauthors = Semino O, Passarino G, Oefner PJ, Lin AA, Arbuzova S, Beckman LE, De Benedictis G, Francalacci P, Kouvatsi A, Limborska S, Marcikiae M, Mika A, Mika B, Primorac D, Santachiara-Benerecetti AS, Cavalli-Sforza LL, Underhill PA | title = The genetic legacy of Paleolithic Homo sapiens sapiens in extant Europeans: a Y chromosome perspective | journal = Science | volume = 290 | issue = 5494 | pages = 1155–9 | date = November 2000 | pmid = 11073453 | doi = 10.1126/science.290.5494.1155 | display-authors = 6 | bibcode = 2000Sci...290.1155S}} but by about 2010 it was thought to have formed near the beginning of the Neolithic Revolution, about 10,000 years ago.[http://www.isogg.org/tree/ISOGG_HapgrpR.html International Society of Genetic Genealogy (ISOGG) – Y-DNA Haplogroup R and its Subclades]{{Cite book| vauthors = Arredi B, Poloni ES, Tyler-Smith C |chapter=The peopling of Europe |editor=Crawford, Michael H. |title=Anthropological genetics: theory, methods and applications |url=https://archive.org/details/anthropologicalg00craw |url-access=limited |publisher=Cambridge University Press |location=Cambridge, UK |year=2007 |page=[https://archive.org/details/anthropologicalg00craw/page/n404 394] |isbn=978-0-521-54697-3}}{{cite journal |vauthors = Cruciani F, Trombetta B, Antonelli C, Pascone R, Valesini G, Scalzi V, Vona G, Melegh B, Zagradisnik B, Assum G, Efremov GD, Sellitto D, Scozzari R | title = Strong intra- and inter-continental differentiation revealed by Y chromosome SNPs M269, U106 and U152 |journal = Forensic Science International. Genetics |volume = 5 |issue = 3 |pages = e49–52 |date = June 2011 | pmid = 20732840 |doi = 10.1016/j.fsigen.2010.07.006 |display-authors=6}} More recent archaeogenetics studies since 2015, however, strongly suggest an origin among Eneolithic hunter-gatherers from eastern Europe.{{cite journal | vauthors = Allentoft ME, Sikora M, Sjögren KG, Rasmussen S, Rasmussen M, Stenderup J, Damgaard PB, Schroeder H, Ahlström T, Vinner L, Malaspinas AS, Margaryan A, Higham T, Chivall D, Lynnerup N, Harvig L, Baron J, Della Casa P, Dąbrowski P, Duffy PR, Ebel AV, Epimakhov A, Frei K, Furmanek M, Gralak T, Gromov A, Gronkiewicz S, Grupe G, Hajdu T, Jarysz R, Khartanovich V, Khokhlov A, Kiss V, Kolář J, Kriiska A, Lasak I, Longhi C, McGlynn G, Merkevicius A, Merkyte I, Metspalu M, Mkrtchyan R, Moiseyev V, Paja L, Pálfi G, Pokutta D, Pospieszny Ł, Price TD, Saag L, Sablin M, Shishlina N, Smrčka V, Soenov VI, Szeverényi V, Tóth G, Trifanova SV, Varul L, Vicze M, Yepiskoposyan L, Zhitenev V, Orlando L, Sicheritz-Pontén T, Brunak S, Nielsen R, Kristiansen K, Willerslev E | display-authors = 6 | title = Population genomics of Bronze Age Eurasia | journal = Nature | volume = 522 | issue = 7555 | pages = 167–172 | date = June 2015 | pmid = 26062507 | doi = 10.1038/nature14507 | s2cid = 4399103 | bibcode = 2015Natur.522..167A | url = https://depot.ceon.pl/handle/123456789/13155 }}
Balaresque et al. (2010) based on the pattern of Y-STR diversity argued for a single source in the Near East and introduction to Europe via Anatolia in the Neolithic Revolution. In this scenario, Mesolithic hunter-gatherers in Europe would have been nearly replaced by the incoming farmers. By contrast, Busby et al. (2012) could not confirm the results of Balaresque et al. (2010) and could not make credible estimates of the age of R-M269 based on Y-STR diversity.{{cite journal | vauthors = Busby GB, Brisighelli F, Sánchez-Diz P, Ramos-Luis E, Martinez-Cadenas C, Thomas MG, Bradley DG, Gusmão L, Winney B, Bodmer W, Vennemann M, Coia V, Scarnicci F, Tofanelli S, Vona G, Ploski R, Vecchiotti C, Zemunik T, Rudan I, Karachanak S, Toncheva D, Anagnostou P, Ferri G, Rapone C, Hervig T, Moen T, Wilson JF, Capelli C | display-authors = 6 | title = The peopling of Europe and the cautionary tale of Y chromosome lineage R-M269 | journal = Proceedings. Biological Sciences | volume = 279 | issue = 1730 | pages = 884–892 | date = March 2012 | pmid = 21865258 | pmc = 3259916 | doi = 10.1098/rspb.2011.1044 }} Furthermore, more recent studies have found that the Y-DNA of Early European Farmers is typically haplogroup G2a.{{cite journal | vauthors = Mathieson I, Lazaridis I, Rohland N, Mallick S, Patterson N, Roodenberg SA, Harney E, Stewardson K, Fernandes D, Novak M, Sirak K, Gamba C, Jones ER, Llamas B, Dryomov S, Pickrell J, Arsuaga JL, de Castro JM, Carbonell E, Gerritsen F, Khokhlov A, Kuznetsov P, Lozano M, Meller H, Mochalov O, Moiseyev V, Guerra MA, Roodenberg J, Vergès JM, Krause J, Cooper A, Alt KW, Brown D, Anthony D, Lalueza-Fox C, Haak W, Pinhasi R, Reich D | display-authors = 6 | title = Genome-wide patterns of selection in 230 ancient Eurasians | journal = Nature | volume = 528 | issue = 7583 | pages = 499–503 | date = December 2015 | pmid = 26595274 | pmc = 4918750 | doi = 10.1038/nature16152 | bibcode = 2015Natur.528..499M }}
According to a 2015 study, a hunter-gatherer from Samara (dated 5640-5555 cal BCE) belonging to haplogroup R1b1(*) was ancestral for both haplogroups R-M269 and R-M478. According to the authors, the occurrence of basal forms of R1b in eastern European hunter-gatherers provides a "geographically plausible source" for haplogroup R-M269. Subclades of R-M269, such as R-Z2103, have been found to be prevalent in ancient DNA found in individuals associated with the Yamnaya culture and related populations,{{cite journal | vauthors = Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, Llamas B, Brandt G, Nordenfelt S, Harney E, Stewardson K, Fu Q, Mittnik A, Bánffy E, Economou C, Francken M, Friederich S, Pena RG, Hallgren F, Khartanovich V, Khokhlov A, Kunst M, Kuznetsov P, Meller H, Mochalov O, Moiseyev V, Nicklisch N, Pichler SL, Risch R, Rojo Guerra MA, Roth C, Szécsényi-Nagy A, Wahl J, Meyer M, Krause J, Brown D, Anthony D, Cooper A, Alt KW, Reich D | display-authors = 6 | title = Massive migration from the steppe was a source for Indo-European languages in Europe | journal = Nature | volume = 522 | issue = 7555 | pages = 207–11 | date = June 2015 | pmid = 25731166 | pmc = 5048219 | doi = 10.1038/nature14317 | arxiv = 1502.02783 | bibcode = 2015Natur.522..207H }}{{cite journal | vauthors = Olalde I, Brace S, Allentoft ME, Armit I, Kristiansen K, Booth T, Rohland N, Mallick S, Szécsényi-Nagy A, Mittnik A, Altena E, Lipson M, Lazaridis I, Harper TK, Patterson N, Broomandkhoshbacht N, Diekmann Y, Faltyskova Z, Fernandes D, Ferry M, Harney E, de Knijff P, Michel M, Oppenheimer J, Stewardson K, Barclay A, Alt KW, Liesau C, Ríos P, Blasco C, Miguel JV, García RM, Fernández AA, Bánffy E, Bernabò-Brea M, Billoin D, Bonsall C, Bonsall L, Allen T, Büster L, Carver S, Navarro LC, Craig OE, Cook GT, Cunliffe B, Denaire A, Dinwiddy KE, Dodwell N, Ernée M, Evans C, Kuchařík M, Farré JF, Fowler C, Gazenbeek M, Pena RG, Haber-Uriarte M, Haduch E, Hey G, Jowett N, Knowles T, Massy K, Pfrengle S, Lefranc P, Lemercier O, Lefebvre A, Martínez CH, Olmo VG, Ramírez AB, Maurandi JL, Majó T, McKinley JI, McSweeney K, Mende BG, Modi A, Kulcsár G, Kiss V, Czene A, Patay R, Endrődi A, Köhler K, Hajdu T, Szeniczey T, Dani J, Bernert Z, Hoole M, Cheronet O, Keating D, Velemínský P, Dobeš M, Candilio F, Brown F, Fernández RF, Herrero-Corral AM, Tusa S, Carnieri E, Lentini L, Valenti A, Zanini A, Waddington C, Delibes G, Guerra-Doce E, Neil B, Brittain M, Luke M, Mortimer R, Desideri J, Besse M, Brücken G, Furmanek M, Hałuszko A, Mackiewicz M, Rapiński A, Leach S, Soriano I, Lillios KT, Cardoso JL, Pearson MP, Włodarczak P, Price TD, Prieto P, Rey PJ, Risch R, Rojo Guerra MA, Schmitt A, Serralongue J, Silva AM, Smrčka V, Vergnaud L, Zilhão J, Caramelli D, Higham T, Thomas MG, Kennett DJ, Fokkens H, Heyd V, Sheridan A, Sjögren KG, Stockhammer PW, Krause J, Pinhasi R, Haak W, Barnes I, Lalueza-Fox C, Reich D | display-authors = 6 | title = The Beaker phenomenon and the genomic transformation of northwest Europe | journal = Nature | volume = 555 | issue = 7695 | pages = 190–196 | date = March 2018 | pmid = 29466337 | pmc = 5973796 | doi = 10.1038/nature25738 | bibcode = 2018Natur.555..190O }} and the dispersal of this haplogroup is associated with the spread of so-called "steppe ancestry" and at least some of the Indo-European languages.{{cite book | vauthors = Anthony DW | chapter =Ancient DNA, Mating Networks, and the Anatolian Split|date=2019-12-06| doi = 10.1163/9789004416192_003 | veditors = Serangeli M, Olander T | title =Dispersals and Diversification|pages=21–53|publisher=BRILL|isbn=978-90-04-41619-2 | s2cid =213909442}}
According to Lazaridis et al. (2022), "the most likely hypothesis" is that the entire R-M269 clade originated "in the North Caucasus and steppe to the north".{{cite journal |last1=Lazaridis |first1=Iosif |display-authors=etal |date=2022 |title=The genetic history of the Southern Arc: a bridge between West Asia and Europe |journal=Science |volume=377 |issue=6609 |pages=Supplementary material, p.332 |doi=10.1126/science.abm4247 |pmid=36007055 |pmc=10064553 |quote=Given that within the phylogeny of R-M269 (R-PF7562, (R-L51, R-Z2103 is meant) both R-PF7562 and R-Z2103 have their earliest examples in the North Caucasus and steppe to the north, the most likely hypothesis is that the entire R-M269 clade originated there as well, with R-L51 representing a lineage that eventually became highly successful in mainland Europe, R-PF7562 a lineage that did not achieve the prominence of its relatives, and R-Z2103 became highly successful (briefly) as part of the Yamnaya culture and its offshoots}}
The subclade R-P311 is substantially confined to Western Europe in modern populations. R-P311 is absent from Neolithic-era ancient DNA found in Western Europe, strongly suggesting that its current distribution is due to population movements within Europe taking place after the end of the Neolithic. The three major subclades of P311 are U106 (S21), L21 (M529, S145), and U152 (S28). These show a clear articulation within Western Europe, with centers in the Low Countries, the British Isles and the Alps, respectively.{{cite conference | vauthors = Hammer M | title = Origins of R-M269 Diversity in Europe | conference = FamilyTreeDNA 9th Annual Conference | year = 2013 }} These lineages are associated with the non-Iberian steppe-related groups of the Bell Beaker culture, and demonstrate the relationship between steppe-related ancestry and R1b-M269 subclades, which are "the major lineage associated with the arrival of Steppe ancestry in western Europe after 2500 BC".{{cite journal | vauthors = Sjögren KG, Olalde I, Carver S, Allentoft ME, Knowles T, Kroonen G, Pike AW, Schröter P, Brown KA, Brown KR, Harrison RJ, Bertemes F, Reich D, Kristiansen K, Heyd V | display-authors = 6 | title = Kinship and social organization in Copper Age Europe. A cross-disciplinary analysis of archaeology, DNA, isotopes, and anthropology from two Bell Beaker cemeteries | journal = PLOS ONE | volume = 15 | issue = 11 | pages = e0241278 | date = 2019-12-11 | pmid = 33196640 | doi = 10.1371/journal.pone.0241278| doi-access = free |biorxiv=10.1101/863944 | pmc = 7668604 | hdl-access = free | s2cid = 212833639 | hdl = 10261/236801}}
Distribution
{{Disputed|date=August 2020}}
European R1b is dominated by R-M269. It has been found at generally low frequencies throughout central Eurasia,{{cite journal | vauthors = Underhill PA, Shen P, Lin AA, Jin L, Passarino G, Yang WH, Kauffman E, Bonné-Tamir B, Bertranpetit J, Francalacci P, Ibrahim M, Jenkins T, Kidd JR, Mehdi SQ, Seielstad MT, Wells RS, Piazza A, Davis RW, Feldman MW, Cavalli-Sforza LL, Oefner PJ | display-authors = 6 | title = Y chromosome sequence variation and the history of human populations | journal = Nature Genetics | volume = 26 | issue = 3 | pages = 358–61 | date = November 2000 | pmid = 11062480 | doi = 10.1038/81685 | s2cid = 12893406}} but with relatively high frequency among the Bashkirs of the Perm region (84.0%) and Baymaksky District (81.0%).{{cite web | url = http://ftp.anrb.ru/molgen/Lobov_AS.PDF | vauthors = Lobov AS, etal | year = 2009 | title = Structure of the Gene Pool of Bashkir Subpopulations | language = ru | archive-url = https://web.archive.org/web/20110816193639/http://ftp.anrb.ru/molgen/Lobov_AS.PDF | archive-date = 2011-08-16 }} This marker is present in China and India at frequencies of less than one percent. The table below lists in more detail the frequencies of M269 in regions in Asia, Europe, and Africa.
Distribution of R-M269 in Europe increases in frequency from east to west. It peaks at the national level in Wales at a rate of 92%, at 82% in Ireland, 70% in Scotland, 68% in Spain, 60% in France (76% in Normandy), about 60% in Portugal,{{cite journal | vauthors = Myres NM, Rootsi S, Lin AA, Järve M, King RJ, Kutuev I, Cabrera VM, Khusnutdinova EK, Pshenichnov A, Yunusbayev B, Balanovsky O, Balanovska E, Rudan P, Baldovic M, Herrera RJ, Chiaroni J, Di Cristofaro J, Villems R, Kivisild T, Underhill PA | display-authors = 6 | title = A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe | journal = European Journal of Human Genetics | volume = 19 | issue = 1 | pages = 95–101 | date = January 2011 | pmid = 20736979 | pmc = 3039512 | doi = 10.1038/ejhg.2010.146 }} 50% in Germany, 50% in the Netherlands, 47% in Italy,{{cite journal | vauthors = Grugni V, Raveane A, Mattioli F, Battaglia V, Sala C, Toniolo D, Ferretti L, Gardella R, Achilli A, Olivieri A, Torroni A, Passarino G, Semino O | display-authors = 6 | title = Reconstructing the genetic history of Italians: new insights from a male (Y-chromosome) perspective | journal = Annals of Human Biology | volume = 45 | issue = 1 | pages = 44–56 | date = February 2018 | pmid = 29382284 | doi = 10.1080/03014460.2017.1409801 | s2cid = 43501209 }} 45% in Eastern England, 43% in Denmark and 42% in Iceland. It is as high as 95% in parts of Ireland. It is also found in some areas of North Africa, where its frequency peaks at 10% in some parts of Algeria.{{cite journal | vauthors = Robino C, Crobu F, Di Gaetano C, Bekada A, Benhamamouch S, Cerutti N, Piazza A, Inturri S, Torre C | display-authors = 6 | title = Analysis of Y-chromosomal SNP haplogroups and STR haplotypes in an Algerian population sample | journal = International Journal of Legal Medicine | volume = 122 | issue = 3 | pages = 251–255 | date = May 2008 | pmid = 17909833 | doi = 10.1007/s00414-007-0203-5 | s2cid = 11556974 }} M269 has likewise been observed among 8% of the Herero in Namibia.{{cite journal | vauthors = Wood ET, Stover DA, Ehret C, Destro-Bisol G, Spedini G, McLeod H, Louie L, Bamshad M, Strassmann BI, Soodyall H, Hammer MF | display-authors = 6 | title = Contrasting patterns of Y chromosome and mtDNA variation in Africa: evidence for sex-biased demographic processes | journal = European Journal of Human Genetics | volume = 13 | issue = 7 | pages = 867–876 | date = July 2005 | pmid = 15856073 | doi = 10.1038/sj.ejhg.5201408 | s2cid = 20279122 | doi-access = free }} The R-M269 subclade has been found in ancient Guanche (Bimbapes) fossils excavated in Punta Azul, El Hierro, Canary Islands, which are dated to the 10th century (~44%).{{cite journal|vauthors=Ordóñez AC, Fregel R, Trujillo-Mederos A, Hervella M, de-la-Rúa C, Arnay-de-la-Rosa M |title=Genetic studies on the prehispanic population buried in Punta Azul cave (El Hierro, Canary Islands)|journal=Journal of Archaeological Science|date=2017|volume=78|pages=20–28|doi=10.1016/j.jas.2016.11.004|bibcode=2017JArSc..78...20O }} In western Asia, R-M269 has been reported in 29.2% of Assyrian males from Iran.{{cite journal | vauthors = Grugni V, Battaglia V, Hooshiar Kashani B, Parolo S, Al-Zahery N, Achilli A, Olivieri A, Gandini F, Houshmand M, Sanati MH, Torroni A, Semino O | display-authors = 6 | title = Ancient migratory events in the Middle East: new clues from the Y-chromosome variation of modern Iranians | journal = PLOS ONE | volume = 7 | issue = 7 | pages = e41252 | year = 2012 | pmid = 22815981 | pmc = 3399854 | doi = 10.1371/journal.pone.0041252 | doi-access = free | bibcode = 2012PLoSO...741252G }} Haplogroup R1b1 and its subclades in Asia.Proceedings of the Russian Academy of DNA Genealogy, 3, 1676–1695 (in Russian).
M269* (xL23) is found at highest frequency in the central Balkans notably Kosovo with 7.9%, North Macedonia 5.1% and Serbia 4.4%. Kosovo is notable in having a high percentage of descendant L23* or L23(xM412) at 11.4% unlike most other areas with significant percentages of M269* and L23* except for Poland with 2.4% and 9.5% and the Bashkirs of southeast Bashkortostan with 2.4% and 32.2% respectively. Notably this Bashkir population also has a high percentage of M269 sister branch M73 at 23.4%. Five individuals out of 110 tested in the Ararat Valley, Armenia belonged to R1b1a2* and 36 to L23*, with none belonging to known subclades of L23. Trofimova et al. (2015) found a surprising high frequency of R1b-L23 (Z2105/2103) among the peoples of the Idel-Ural. 21 out of 58 (36.2%) of Burzyansky District Bashkirs, 11 out of 52 (21.2%) of Udmurts, 4 out of 50 (8%) of Komi, 4 out of 59 (6.8%) of Mordvins, 2 out of 53 (3.8%) of Besermyan and 1 out of 43 (2.3%) of Chuvash were R1b-L23 (Z2105/2103),Трофимова Натал'я Вадимовна (Feb. 2015), [http://ibg.anrb.ru/disovet/zashita/2015/02Trofimova/2015_02_TrofimovaAvtoref.pdf "Изменчивость Митохондриальной ДНК и Y-Хромосомы в Популяциях Волго-Уральского Региона"] {{Webarchive|url=https://web.archive.org/web/20170402192825/http://ibg.anrb.ru/disovet/zashita/2015/02Trofimova/2015_02_TrofimovaAvtoref.pdf |date=2017-04-02 }} ("Mitochondrial DNA variation and the Y-chromosome in the population of the Volga-Ural Region"). Автореферат. диссертации на соискание ученой степени кандидата биологических наук. Уфа – 2015. the type of R1b found in the recently analyzed Yamna remains of the Samara Oblast and Orenburg Oblast.{{cite journal | vauthors = Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, Llamas B, Brandt G, Nordenfelt S, Harney E, Stewardson K, Fu Q, Mittnik A, Bánffy E, Economou C, Francken M, Friederich S, Pena RG, Hallgren F, Khartanovich V, Khokhlov A, Kunst M, Kuznetsov P, Meller H, Mochalov O, Moiseyev V, Nicklisch N, Pichler SL, Risch R, Rojo Guerra MA, Roth C, Szécsényi-Nagy A, Wahl J, Meyer M, Krause J, Brown D, Anthony D, Cooper A, Alt KW, Reich D | display-authors = 6 | title = Massive migration from the steppe was a source for Indo-European languages in Europe | journal = Nature | volume = 522 | issue = 7555 | pages = 207–11 | date = June 2015 | pmid = 25731166 | doi = 10.1038/nature14317 | name-list-style = vanc | biorxiv = 10.1101/013433 | pmc = 5048219 | arxiv = 1502.02783 | bibcode = 2015Natur.522..207H }}
Especially Western European R1b is dominated by specific sub-clades of R-M269 (with some small amounts of other types found in areas such as Sardinia{{cite journal | vauthors = Morelli L, Contu D, Santoni F, Whalen MB, Francalacci P, Cucca F | title = A comparison of Y-chromosome variation in Sardinia and Anatolia is more consistent with cultural rather than demic diffusion of agriculture | journal = PLOS ONE | volume = 5 | issue = 4 | pages = e10419 | date = April 2010 | pmid = 20454687 | pmc = 2861676 | doi = 10.1371/journal.pone.0010419 | veditors = Lalueza-Fox C | bibcode = 2010PLoSO...510419M | doi-access = free }}).
Within Europe, R-M269 is dominated by R-M412, also known as R-L51, which according to Myres et al. (2010) is "virtually absent in the Near East, the Caucasus and West Asia." This Western European population is further divided between R-P312/S116 and R-U106/S21, which appear to spread from the western and eastern Rhine river basin respectively.
Myres et al. note further that concerning its closest relatives, in R-L23*, it is "instructive" that these are often more than 10% of the population in the Caucasus, Turkey, and some southeast European and circum-Uralic populations.
In Western Europe it is present but in generally much lower levels apart from "an instance of 27% in Switzerland's Upper Rhone Valley."
In addition, the sub-clade distribution map, Figure 1h titled "L11(xU106,S116)", in Myres et al. shows that R-P310/L11* (or as yet undefined subclades of R-P310/L11) occurs only in frequencies greater than 10% in Central England with surrounding areas of England and Wales having lower frequencies. This R-P310/L11* is almost non-existent in the rest of Eurasia and North Africa with the exception of coastal lands fringing the western and southern Baltic (reaching 10% in Eastern Denmark and 6% in northern Poland) and in Eastern Switzerland and surrounds.
{{Clade
| label1=M269 (R1b1a1a2)ISOGG tree as of 2017 ([https://isogg.org/tree/ISOGG_HapgrpR.html isogg.org])
| 1={{Clade
| label1=
| 1= R-M269*
| label2=L23 (R1b1a1a2a)
| 2={{Clade
| label1=
| 1= R-L23*: Caucasus, Turkey, circum-Uralic; Upper Rhone Valley
| label3=Z2103 (R1b1a1a2a2)
| 3=Z2103: Balkans and Turkey, Samara (Russia, Yamnaya a.c.), South Ural (burjan bashkirs)
| label2=L51/M412 (R1b1a1a2a1)
| 2={{Clade
| label1=
| 1=R-L51*/R-M412*: Central France
| label2=L151/P310/P311 (R1b1a1a2a1a)
| 2={{Clade
| label1=
| 1=R-P310/L11*: Central England
| label2=U106 (R1b1a1a2a1a1)
| 2= R-U106: Netherlands, England, Norway; Germanic Europe
| label3=P312/S116 (R1b1a1a2a1a2)
| 3={{Clade
| label1=
| 1=S116*: Iberian Peninsula
| label2=U152 (R1b1a1a2a1a2b)
|2 =U152: Corsica, Sardinia; Northern Italy, Central Italy, Switzerland, Central France, Russia (Perm region, Ghaeynae bashkirs)
|label3=L21_M529_S145 (R1b1a1a2a1a2c1)
|3 = M529: Brittany, Ireland, Scotland, Wales}}
|label4=CTS4528 (R1b1a1a2a1a3a)
|4=R-CTS4528 }}}}
}}
}}
}}
In 2009, DNA extracted from the femur bones of 6 skeletons in an early-medieval burial place in Ergolding (Bavaria, Germany) dated to around c. 670 yielded the following results: 4 were found to be haplogroup R1b with the closest matches in modern populations of Germany, Ireland and the USA while 2 were in Haplogroup G2a.{{cite journal | vauthors = Vanek D, Saskova L, Koch H | title = Kinship and Y-chromosome analysis of 7th century human remains: novel DNA extraction and typing procedure for ancient material | journal = Croatian Medical Journal | volume = 50 | issue = 3 | pages = 286–95 | date = June 2009 | pmid = 19480023 | pmc = 2702742 | doi = 10.3325/cmj.2009.50.286 | series = 3 }}
Population studies which test for M269 have become more common in recent years, while in earlier studies men in this haplogroup are only visible in the data by extrapolation of what is likely. The following gives a summary of most of the studies which specifically tested for M269, showing its distribution (as a percentage of total population) in Europe, North Africa, the Middle East and Central Asia as far as China and Nepal.
Sub-clades
{{Disputed|date=August 2020}}
=R1b1a1a2a (R-L23) =
R-L23* (R1b1a1a2a*) is now most commonly found in Europe, Anatolia, the Caucasus.
= R1b1a1a2a1 (R-L51) =
= R1b1a1a2a1a (R-L151) =
{{main|Haplogroup R-L151}}
R-L151 (L151/PF6542, CTS7650/FGC44/PF6544/S1164, L11, L52/PF6541, P310/PF6546/S129, P311/PF6545/S128) also known as R1b1a1a2a1, and its subclades, include most males with R1b in Western Europe.
= R1b1a1a2a1a1 (R-U106) =
This subclade is defined by the presence of the SNP U106, also known as S21 and M405.{{cite journal | vauthors = Sims LM, Garvey D, Ballantyne J | title = Sub-populations within the major European and African derived haplogroups R1b3 and E3a are differentiated by previously phylogenetically undefined Y-SNPs | journal = Human Mutation | volume = 28 | issue = 1 | pages = 97 | date = January 2007 | pmid = 17154278 | doi = 10.1002/humu.9469 | s2cid = 34556775 | doi-access = free }} It appears to represent over 25% of R1b in Europe. In terms of percentage of total population, its epicenter is Friesland, where it makes up 44% of the population.{{cite web | title = Origins of R-M269 Diversity in Europe | url = https://gap.familytreedna.com/media/docs/2013/Hammer_M269_Diversity_in_Europe.pdf }} In terms of total population numbers, its epicenter is Central Europe, where it comprises 60% of R1 combined. See also Haplogroup R-Z18
{{Clade
| label1=U106/S21/M405
| 1={{Clade
| label1=un-defined
| 1=R-U106* (R-U106-*)
| label2=FGC3861
| 2=R-FGC3861 (R1b1a2a1a1a)
| label3=Z18
| 3=R-Z18 (R1b1a2a1a1b)
| label4=Z381
| 4={{Clade
| label1=S264
| 1=R-S264 (R1b1a2a1a1c1)
| label2=S499
| 2=R-S499 (R1b1a2a1a1c2)
| label3=M1994
| 3=R-M1994 (R1b1a2a1a1c3)}}
| label5=FGC396
| 5=R-FGC396 (R1b1a2a1a1d)
| label6=S12025
|6=R-S12025 (R1b1a2a1a1e)}}
}}
While this sub-clade of R1b is frequently discussed amongst genetic genealogists, the following table represents the peer-reviewed findings published so far in the 2007 articles of Myres et al. and Sims et al.{{cite journal | vauthors = Myres NM, Ekins JE, Lin AA, Cavalli-Sforza LL, Woodward SR, Underhill PA | title = Y-chromosome short tandem repeat DYS458.2 non-consensus alleles occur independently in both binary haplogroups J1-M267 and R1b3-M405 | journal = Croatian Medical Journal | volume = 48 | issue = 4 | pages = 450–9 | date = August 2007 | pmid = 17696299 | pmc = 2080563 | url = http://www.cmj.hr/2007/48/4/17696299.htm }}
= R-P312 =
R1b1a1a2a1a2, better known as R-P312 (or R-S116) is one of the most common types of R-M269 in Europe, alongside R-U106. Myres et al. described it as originating in and spreading from the west of the Rhine basin.
R-P312 has been the subject of significant, ongoing study concerning its complex internal structure.
{{Clade
| label1=P312
| 1={{Clade
| label1=
| 1=R-P312*
| label2=DF27
| 2={{Clade
|1=R-S227/Z196
|2=R-Z2552
|3=R-L881
|4=R-A431 }}
| label3=U152
| 3={{Clade
| 1=R-L2
| 2=R-S206
| 3=R-Z56 }}
| label4=L21
| 4={{Clade
| 1=R-A7905
| 2=R-A5846
| 3=R-DF63 (R-S522)
| 4=R-DF13 (R-CTS241/R-S521) }}
| 5=R-L238
| 6=R-DF19
| 7=R-DF99 }}}}
==R-DF27==
{{main|Haplogroup R-DF27}}
R-M153
R-M153 is a subclade of R-DF27 that has been found mostly in Basques and Gascons, among whom it represents a sizeable fraction of the Y-DNA pool,{{cite journal | vauthors = Adams SM, Bosch E, Balaresque PL, Ballereau SJ, Lee AC, Arroyo E, López-Parra AM, Aler M, Grifo MS, Brion M, Carracedo A, Lavinha J, Martínez-Jarreta B, Quintana-Murci L, Picornell A, Ramon M, Skorecki K, Behar DM, Calafell F, Jobling MA | title = The genetic legacy of religious diversity and intolerance: paternal lineages of Christians, Jews, and Muslims in the Iberian Peninsula | journal = American Journal of Human Genetics | volume = 83 | issue = 6 | pages = 725–36 | date = December 2008 | pmid = 19061982 | pmc = 2668061 | doi = 10.1016/j.ajhg.2008.11.007 | display-authors = 6 }}{{cite journal | vauthors = López-Parra AM, Gusmão L, Tavares L, Baeza C, Amorim A, Mesa MS, Prata MJ, Arroyo-Pardo E | title = In search of the pre- and post-neolithic genetic substrates in Iberia: evidence from Y-chromosome in Pyrenean populations | journal = Annals of Human Genetics | volume = 73 | issue = 1 | pages = 42–53 | date = January 2009 | pmid = 18803634 | doi = 10.1111/j.1469-1809.2008.00478.x | s2cid = 43273988 }} though is also found occasionally among Iberians in general. The first time it was located (Bosch 2001{{cite journal | vauthors = Bosch E, Calafell F, Comas D, Oefner PJ, Underhill PA, Bertranpetit J | title = High-resolution analysis of human Y-chromosome variation shows a sharp discontinuity and limited gene flow between northwestern Africa and the Iberian Peninsula | journal = American Journal of Human Genetics | volume = 68 | issue = 4 | pages = 1019–29 | date = April 2001 | pmid = 11254456 | pmc = 1275654 | doi = 10.1086/319521 }}) it was described as H102 and included seven Basques and one Andalusian.
R-M167 is a subclade of R-DF27 defined by the presence of the marker M167. The first author to test for this marker (long before current haplogroup nomenclature existed) was Hurles in 1999, who tested 1158 men in various populations.{{cite journal | vauthors = Hurles ME, Veitia R, Arroyo E, Armenteros M, Bertranpetit J, Pérez-Lezaun A, Bosch E, Shlumukova M, Cambon-Thomsen A, McElreavey K, López De Munain A, Röhl A, Wilson IJ, Singh L, Pandya A, Santos FR, Tyler-Smith C, Jobling MA | title = Recent male-mediated gene flow over a linguistic barrier in Iberia, suggested by analysis of a Y-chromosomal DNA polymorphism | journal = American Journal of Human Genetics | volume = 65 | issue = 5 | pages = 1437–48 | date = November 1999 | pmid = 10521311 | pmc = 1288297 | doi = 10.1086/302617 | display-authors = 6 }} He found it relatively common among Basques (13/117: 11%) and Catalans (7/32: 22%). Other occurrences were found among other French, British, Spaniards, Béarnais, and Germans.
In 2000 Rosser et al., in a study which tested 3616 men in various populations{{cite journal | vauthors = Rosser ZH, Zerjal T, Hurles ME, Adojaan M, Alavantic D, Amorim A, Amos W, Armenteros M, Arroyo E, Barbujani G, Beckman G, Beckman L, Bertranpetit J, Bosch E, Bradley DG, Brede G, Cooper G, Côrte-Real HB, de Knijff P, Decorte R, Dubrova YE, Evgrafov O, Gilissen A, Glisic S, Gölge M, Hill EW, Jeziorowska A, Kalaydjieva L, Kayser M, Kivisild T, Kravchenko SA, Krumina A, Kucinskas V, Lavinha J, Livshits LA, Malaspina P, Maria S, McElreavey K, Meitinger TA, Mikelsaar AV, Mitchell RJ, Nafa K, Nicholson J, Nørby S, Pandya A, Parik J, Patsalis PC, Pereira L, Peterlin B, Pielberg G, Prata MJ, Previderé C, Roewer L, Rootsi S, Rubinsztein DC, Saillard J, Santos FR, Stefanescu G, Sykes BC, Tolun A, Villems R, Tyler-Smith C, Jobling MA | title = Y-chromosomal diversity in Europe is clinal and influenced primarily by geography, rather than by language | journal = American Journal of Human Genetics | volume = 67 | issue = 6 | pages = 1526–43 | date = December 2000 | pmid = 11078479 | pmc = 1287948 | doi = 10.1086/316890 | display-authors = 6 }} also tested for that same marker, naming the haplogroup Hg22, and again it was found mainly among Basques (19%), in lower frequencies among French (5%), Bavarians (3%), Spaniards (2%), Southern Portuguese (2%), and in single occurrences among Romanians, Slovenians, Dutch, Belgians and English.::In 2001 Bosch described this marker as H103, in 5 Basques and 5 Catalans. Further regional studies have located it in significant amounts in Asturias, Cantabria and Galicia, as well as again among Basques. Cases in the Azores have been reported.{{Citation needed|date=May 2011}} In 2008 two research papers by López-Parra and Adams, respectively, confirmed a strong association with all or most of the Pyrenees and Eastern Iberia.
In a larger study of Portugal in 2006, with 657 men tested, Beleza et al. confirmed similar low levels in all the major regions, from 1.5%–3.5%.{{cite journal | vauthors = Beleza S, Gusmão L, Lopes A, Alves C, Gomes I, Giouzeli M, Calafell F, Carracedo A, Amorim A | title = Micro-phylogeographic and demographic history of Portuguese male lineages | journal = Annals of Human Genetics | volume = 70 | issue = Pt 2 | pages = 181–94 | date = March 2006 | pmid = 16626329 | doi = 10.1111/j.1529-8817.2005.00221.x | s2cid = 4652154 | quote = 395/657 | display-authors = 6 }}
R-L165
This subclade is defined by the presence of the marker S68, also known as L165. It is found in England, Scandinavia, and Scotland (in this country it is mostly found in the Northern Isles and Outer Hebrides). It has been suggested, therefore, that it arrived in the British Isles with Vikings.{{Cite book | vauthors = Moffat A, Wilson JF |author1-link=Alistair Moffat |title=The Scots: a genetic journey |url= https://archive.org/details/scotsgeneticjour0000moff |url-access=registration |publisher=Birlinn |year=2011 |pages=[https://archive.org/details/scotsgeneticjour0000moff/page/181 181]–182, 192 |isbn=978-0-85790-020-3 }}
==R-U152==
{{anchor|R-U152}}
R-U152 is defined by the presence of the marker U152, also called S28. Its existence was confirmed by Sims et al. (2007). Myres et al. report this clade "is most frequent (20–44%) in Switzerland, Italy, France and Western Poland, with additional instances exceeding 15% in some regions of England and Germany." Similarly Cruciani et al. (2010){{cite journal|vauthors=Cruciani F, Trombetta B, Antonelli C, Pascone R, Valesini G, Scalzi V, Vona G, Melegh B, Zagradisnik B, Assum G, Efremov GD, Sellitto D, Scozzari R|date=June 2011|title=Strong intra- and inter-continental differentiation revealed by Y chromosome SNPs M269, U106 and U152|journal=Forensic Science International. Genetics|volume=5|issue=3|pages=e49–52|doi=10.1016/j.fsigen.2010.07.006|pmid=20732840}} reported frequency peaks in Northern and Central Italy and France. Out of a sample of 135 men in Tyrol, Austria, 9 tested positive for U152/S28.{{Cite journal | vauthors = Niederstätter H, Berger B, Erhart D, Parson W |title=Recently introduced Y-SNPs improve the resolution within Y-chromosome haplogroup R1b in a central European population sample (Tyrol, Austria)|doi=10.1016/j.fsigss.2007.10.158| date=August 2008| journal=Forensic Science International: Genetics Supplement Series|volume=1|pages=226–227|doi-access=free}} Far removed from this apparent core area, Myres et al. also mention a sub-population in north Bashkortostan, where 71% of 70 men tested belong to R-U152. They propose this to be the result of an isolated founder effect. King et al. (2014) reported four living descendants of Henry Somerset, 5th Duke of Beaufort in the male line tested positive for U-152.Henry Somerset was in turn descended in the patrilineal line from John of Gaunt (1340–1399), a son of King Edward III (1312–1377). In the context of the analysis of the remains of Richard III, which proved to belong to haplogroup G2, the possibility of a false-paternity event, most likely between Edward III and Henry Somerset, was discussed; possibly confirming rumors to the effect that John of Gaunt was illegitimate (Jonathan Sumption, Divided Houses: The Hundred Years War III, 2009, p. 274).
{{cite journal | vauthors = King TE, Fortes GG, Balaresque P, Thomas MG, Balding D, Maisano Delser P, Neumann R, Parson W, Knapp M, Walsh S, Tonasso L, Holt J, Kayser M, Appleby J, Forster P, Ekserdjian D, Hofreiter M, Schürer K | title = Identification of the remains of King Richard III | journal = Nature Communications | volume = 5 | issue = 5631 | pages = 5631 | date = December 2014 | pmid = 25463651 | pmc = 4268703 | doi = 10.1038/ncomms6631 | bibcode = 2014NatCo...5.5631K }}
"Y-chromosome haplotypes from male-line relatives and the remains do not match, which could be attributed to a false-paternity event occurring in any of the intervening generations." Ancient samples from the central European Bell Beaker, Hallstatt and Tumulus cultures belonged to this subclade.{{cite journal | vauthors = Allentoft ME, Sikora M, Sjögren KG, Rasmussen S, Rasmussen M, Stenderup J, Damgaard PB, Schroeder H, Ahlström T, Vinner L, Malaspinas AS, Margaryan A, Higham T, Chivall D, Lynnerup N, Harvig L, Baron J, Della Casa P, Dąbrowski P, Duffy PR, Ebel AV, Epimakhov A, Frei K, Furmanek M, Gralak T, Gromov A, Gronkiewicz S, Grupe G, Hajdu T, Jarysz R, Khartanovich V, Khokhlov A, Kiss V, Kolář J, Kriiska A, Lasak I, Longhi C, McGlynn G, Merkevicius A, Merkyte I, Metspalu M, Mkrtchyan R, Moiseyev V, Paja L, Pálfi G, Pokutta D, Pospieszny Ł, Price TD, Saag L, Sablin M, Shishlina N, Smrčka V, Soenov VI, Szeverényi V, Tóth G, Trifanova SV, Varul L, Vicze M, Yepiskoposyan L, Zhitenev V, Orlando L, Sicheritz-Pontén T, Brunak S, Nielsen R, Kristiansen K, Willerslev E | display-authors = 6 | title = Population genomics of Bronze Age Eurasia | journal = Nature | volume = 522 | issue = 7555 | pages = 167–72 | date = June 2015 | pmid = 26062507 | doi = 10.1038/nature14507 | s2cid = 4399103 | bibcode = 2015Natur.522..167A | url = https://depot.ceon.pl/handle/123456789/13155 }}{{cite journal | vauthors = Damgaard PB, Marchi N, Rasmussen S, Peyrot M, Renaud G, Korneliussen T, Moreno-Mayar JV, Pedersen MW, Goldberg A, Usmanova E, Baimukhanov N, Loman V, Hedeager L, Pedersen AG, Nielsen K, Afanasiev G, Akmatov K, Aldashev A, Alpaslan A, Baimbetov G, Bazaliiskii VI, Beisenov A, Boldbaatar B, Boldgiv B, Dorzhu C, Ellingvag S, Erdenebaatar D, Dajani R, Dmitriev E, Evdokimov V, Frei KM, Gromov A, Goryachev A, Hakonarson H, Hegay T, Khachatryan Z, Khaskhanov R, Kitov E, Kolbina A, Kubatbek T, Kukushkin A, Kukushkin I, Lau N, Margaryan A, Merkyte I, Mertz IV, Mertz VK, Mijiddorj E, Moiyesev V, Mukhtarova G, Nurmukhanbetov B, Orozbekova Z, Panyushkina I, Pieta K, Smrčka V, Shevnina I, Logvin A, Sjögren KG, Štolcová T, Taravella AM, Tashbaeva K, Tkachev A, Tulegenov T, Voyakin D, Yepiskoposyan L, Undrakhbold S, Varfolomeev V, Weber A, Wilson Sayres MA, Kradin N, Allentoft ME, Orlando L, Nielsen R, Sikora M, Heyer E, Kristiansen K, Willerslev E | display-authors = 6 | title = 137 ancient human genomes from across the Eurasian steppes | journal = Nature | volume = 557 | issue = 7705 | pages = 369–374 | date = May 2018 | pmid = 29743675 | doi = 10.1038/s41586-018-0094-2 | s2cid = 13670282 | bibcode = 2018Natur.557..369D | hdl = 1887/3202709 | hdl-access = free }} Analyzed Iron Age Latins, Etruscans and Alpine Celts, dating between 1000 and 100 BCE, belonged primarily to haplogroup R1b-U152 (including the clades L2, Z56 and Z193).{{cite journal | vauthors = Antonio ML, Gao Z, Moots HM, Lucci M, Candilio F, Sawyer S, Oberreiter V, Calderon D, Devitofranceschi K, Aikens RC, Aneli S, Bartoli F, Bedini A, Cheronet O, Cotter DJ, Fernandes DM, Gasperetti G, Grifoni R, Guidi A, La Pastina F, Loreti E, Manacorda D, Matullo G, Morretta S, Nava A, Fiocchi Nicolai V, Nomi F, Pavolini C, Pentiricci M, Pergola P, Piranomonte M, Schmidt R, Spinola G, Sperduti A, Rubini M, Bondioli L, Coppa A, Pinhasi R, Pritchard JK | display-authors = 6 | title = Ancient Rome: A genetic crossroads of Europe and the Mediterranean | journal = Science | volume = 366 | issue = 6466 | pages = 708–714 | date = November 2019 | pmid = 31699931 | pmc = 7093155 | doi = 10.1126/science.aay6826 | bibcode = 2019Sci...366..708A }}{{cite journal | vauthors = Posth C, Zaro V, Spyrou MA, Vai S, Gnecchi-Ruscone GA, Modi A, Peltzer A, Mötsch A, Nägele K, Vågene ÅJ, Nelson EA, Radzevičiūtė R, Freund C, Bondioli LM, Cappuccini L, Frenzel H, Pacciani E, Boschin F, Capecchi G, Martini I, Moroni A, Ricci S, Sperduti A, Turchetti MA, Riga A, Zavattaro M, Zifferero A, Heyne HO, Fernández-Domínguez E, Kroonen GJ, McCormick M, Haak W, Lari M, Barbujani G, Bondioli L, Bos KI, Caramelli D, Krause J | display-authors = 6 | title = The origin and legacy of the Etruscans through a 2000-year archeogenomic time transect | language = English | journal = Science Advances | volume = 7 | issue = 39 | pages = eabi7673 | date = September 2021 | pmid = 34559560 | pmc = 8462907 | doi = 10.1126/sciadv.abi7673 | publisher = American Association for the Advancement of Science | bibcode = 2021SciA....7.7673P }} Ancient samples of Cenomani Cisalpine Gauls from Verona who lived between the 3rd and 1st centuries BCE were predominantly R-U152.{{Cite journal |last1=Laffranchi |first1=Zita |last2=Zingale |first2=Stefania |last3=Tecchiati |first3=Umberto |last4=Amato |first4=Alfonsina |last5=Coia |first5=Valentina |last6=Paladin |first6=Alice |last7=Salzani |first7=Luciano |last8=Thompson |first8=Simon R. |last9=Bersani |first9=Marzia |last10=Dori |first10=Irene |last11=Szidat |first11=Sönke |last12=Lösch |first12=Sandra |last13=Ryan-Despraz |first13=Jessica |last14=Arenz |first14=Gabriele |last15=Zink |first15=Albert |date=2024-02-14 |title="Until death do us part". A multidisciplinary study on human- Animal co- burials from the Late Iron Age necropolis of Seminario Vescovile in Verona (Northern Italy, 3rd-1st c. BCE) |journal=PLOS ONE |language=en |volume=19 |issue=2 |pages=e0293434 |doi=10.1371/journal.pone.0293434 |doi-access=free |issn=1932-6203 |pmc=10866530 |pmid=38354185|bibcode=2024PLoSO..1993434L }} U152 is also found at low frequencies of around 3%-4% in both Morocco and Algeria, pointing to direct maritime contacts between the European and North African sides of the western Mediterranean.{{Cite journal |last1=Bekada |first1=Asmahan |last2=Fregel |first2=Rosa |last3=Cabrera |first3=Vicente M. |last4=Larruga |first4=José M. |last5=Pestano |first5=José |last6=Benhamamouch |first6=Soraya |last7=González |first7=Ana M. |date=2013 |title=Introducing the Algerian mitochondrial DNA and Y-chromosome profiles into the North African landscape |journal=PLOS ONE |volume=8 |issue=2 |pages=e56775 |doi=10.1371/journal.pone.0056775 |doi-access=free |issn=1932-6203 |pmc=3576335 |pmid=23431392|bibcode=2013PLoSO...856775B }}
==R-L21==
{{main|Haplogroup R-L21}}
R-L21, also known as R-M529 and R-S145, is most common in Ireland, Scotland and Wales (i.e. at least 25–50% of their male populations. R-L21 has two primary subclades: R-A5846 and R-S552.
R-DF13
A primary subclade of R-S552, R-DF13 – also known as R-S521, R-Z2542 and R-CTS8221 – is one of the most common subclades of R-L21. At least one study estimated that R-DF13 would be found among more than 50% of living Irish males. The following are among the most common subclades within R-DF13.
- R-DF21, a primary subclade of R-DF13, defined by the presence of the marker DF21 a.k.a. S192. R-DF21 makes up about 10% of all L21 men and is circa 3000 years old.{{cite web |url=http://www.familytreedna.com/public/R-DF21/default.aspx |title=R-DF21 and Subclades Project}}
- R-L159.2 is a subclade of R-DF13, defined by the marker L159/S169. It is known as R-L159.2 because of an unrelated parallel mutation (L159.1), found within haplogroup I2a1a1a (a.k.a. I-L158 or I-M26). Consequently, some Y-DNA trees exclude L159/S169 completely, on the basis that it may be an unreliable marker. For instance, as of 2024, Yfull refers to an equivalent subclade as R-FGC80001 (i.e. R-L21 > R-S552 > R-DF13 > R-Z255 > R-FGC80001).[https://www.yfull.com/live/tree/R-FGC80001/ Yfull, 2012–24, R-FGC80001](Access: 9 April 2024.) R-L159.2 appears to be associated with the Laigin, an ethno-tribal group, after whom the Kingdom of Leinster was named. It is common in males from coastal areas surrounding the Irish Sea, including western Wales, the Isle of Man and the Hebrides; R-L159.2 has also been found at significant levels in Norway, western and southern Scottish mainland, parts of England, northwest France, and northern Denmark.{{cite web |url=http://www.familytreedna.com/public/R1b-L159.2/default.aspx?section=goals |title=R-L159 Project Goals }}
- R-L193: this subclade within R-DF13 is defined by the presence of the marker L193. Many surnames with this marker are associated geographically with the western "Border Region" of Scotland. A few other surnames have a Highland association. R-L193 is a relatively young subclade likely born within the last 2000 years.
- R-L226, under R-DF13, is defined by the presence of the marker L226, also known as S168. Commonly referred to as Irish Type III, it is concentrated in central western Ireland and associated with the Dál gCais kindred.{{Cite journal|title=A Set of Distinctive Marker Values Defines a Y-STR Signature for Gaelic Dalcassian Families|url=http://www.jogg.info/51/files/Wright.htm|vauthors=Wright DM|year=2009|journal=Journal of Genetic Genealogy|access-date=2017-03-17|archive-date=2012-08-24|archive-url=https://web.archive.org/web/20120824020357/http://www.jogg.info/51/files/Wright.htm|url-status=dead}}
- R-L371: a subclade within R-DF13 defined by the presence of the marker L371 – sometimes referred to as the "Welsh modal haplotype". It is associated with ancient Welsh kings and princes.{{Cite news|url=https://www.walesonline.co.uk/lifestyle/showbiz/dafydd-iwan-discovers-impressive-celtic-7829517|title=Dafydd Iwan's rare genetic roots unveiled in new project| vauthors = Bevan N |date=2014-09-25|work=walesonline|access-date=2018-04-05}}{{Cite news|url=https://www.dailypost.co.uk/news/north-wales-news/dafydd-iwan-descended-welsh-kings-7838225|title=Dafydd Iwan 'descended from Welsh kings' who ruled in England| vauthors = Bodden T |date=2014-09-26|work=northwales|access-date=2018-04-05}}{{Cite web |url= https://www.familytreedna.com/public/R-17-14-10/ |title=Family Tree DNA - My FamilyTree DNA Project Website Title|website=www.familytreedna.com|access-date=2018-04-05}}