Haplogroup I-M438

{{Short description|Human Y-chromosome DNA haplogroup}} {{Expert needed|Human Genetic History|talk=|reason=Nomenclature of haplogroup(s) and subclades|date=November 2015}} {{Infobox haplogroup | name =I-M438 | map = | origin-date =28–33,000 years ago<ref name="Peter Underhill 2007 pp. 33-42"/> | origin-place = South-Eastern/Eastern Europe | ancestor =I-M170 | descendants =I-L460, I-L1251 | mutations =M438/P215/S31 | members =

I2a1a: Sardinia<ref name=pmid15162323>{{cite journal | vauthors = Rootsi S, Magri C, Kivisild T, Benuzzi G, Help H, Bermisheva M, Kutuev I, Barać L, Pericić M, Balanovsky O, Pshenichnov A, Dion D, Grobei M, Zhivotovsky LA, Battaglia V, Achilli A, Al-Zahery N, Parik J, King R, Cinnioğlu C, Khusnutdinova E, Rudan P, Balanovska E, Scheffrahn W, Simonescu M, Brehm A, Goncalves R, Rosa A, Moisan JP, Chaventre A, Ferak V, Füredi S, Oefner PJ, Shen P, Beckman L, Mikerezi I, Terzić R, Primorac D, Cambon-Thomsen A, Krumina A, Torroni A, Underhill PA, Santachiara-Benerecetti AS, Villems R, Semino O | display-authors = 6 | title = Phylogeography of Y-chromosome haplogroup I reveals distinct domains of prehistoric gene flow in europe | journal = American Journal of Human Genetics | volume = 75 | issue = 1 | pages = 128–137 | date = July 2004 | pmid = 15162323 | pmc = 1181996 | doi = 10.1086/422196 }}</ref> I2a1b: Bosnia and Herzegovina,<ref name=pmid15944443>{{cite journal | vauthors = Pericić M, Lauc LB, Klarić IM, Rootsi S, Janićijevic B, Rudan I, Terzić R, Colak I, Kvesić A, Popović D, Sijacki A, Behluli I, Dordevic D, Efremovska L, Bajec DD, Stefanović BD, Villems R, Rudan P | display-authors = 6 | title = High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations | journal = Molecular Biology and Evolution | volume = 22 | issue = 10 | pages = 1964–1975 | date = October 2005 | pmid = 15944443 | doi = 10.1093/molbev/msi185 | doi-access = free }}</ref> I2a2: Britain, Germany, and Sweden<ref name=pmid15162323/> }}

'''Haplogroup I-M438''', also known as '''I2''' (ISOGG 2019), is a human DNA Y-chromosome haplogroup, a subclade of haplogroup I-M170. Haplogroup I-M438 originated some time around 26,000–31,000 BCE. It originated in Europe and developed into several main subgroups: I2-M438*, I2a-L460, I2b-L415 and I2c-L596.<ref name=pmid15162323/> The haplogroup can be found all over Europe and reaches its maximum frequency in the Dinaric Alps (Balkans) via founder effect, related to the migrations of the Early Slavs to the Balkan peninsula.

The oldest example so far found is that of Hohle Fels (49) from Germany, being at least 14,200 years old.<ref>{{Cite journal |last1=Posth |first1=Cosimo |last2=Yu |first2=He |last3=Ghalichi |first3=Ayshin |last4=Rougier |first4=Hélène |last5=Crevecoeur |first5=Isabelle |last6=Huang |first6=Yilei |last7=Ringbauer |first7=Harald |last8=Rohrlach |first8=Adam B. |last9=Nägele |first9=Kathrin |last10=Villalba-Mouco |first10=Vanessa |last11=Radzeviciute |first11=Rita |last12=Ferraz |first12=Tiago |last13=Stoessel |first13=Alexander |last14=Tukhbatova |first14=Rezeda |last15=Drucker |first15=Dorothée G. |date=2023-03-01 |title=Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers |journal=Nature |language=en |volume=615 |issue=7950 |pages=117–126 |doi=10.1038/s41586-023-05726-0 |pmid=36859578 |pmc=9977688 |bibcode=2023Natur.615..117P |issn=1476-4687|hdl=10256/23099 |hdl-access=free }}</ref>

==Origin and prehistoric presence== {{expand section|date=April 2017}} Haplogroup I2a was the most frequent Y-DNA among Mesolithic Western European hunter-gatherers (WHG) belonging to Villabruna Cluster. A 2015 study found haplogroup I2a in 13,500 year old remains from the Azilian culture (from Grotte du Bichon, modern Switzerland).<ref>{{cite journal | vauthors = 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 | display-authors = 6 | title = Upper Palaeolithic genomes reveal deep roots of modern Eurasians | journal = Nature Communications | volume = 6 | article-number = 8912 | date = November 2015 | pmid = 26567969 | pmc = 4660371 | doi = 10.1038/ncomms9912 | bibcode = 2015NatCo...6.8912J }}</ref> Subclades of I2a1 (I-P37.2), namely I-M423 and I-M26, have been found in remains of WHGs dating from 10,000 to 8,000 years before present.<ref>[http://www.ancestraljourneys.org/mesolithicdna.shtml] {{Webarchive|url=https://web.archive.org/web/20170430223117/http://www.ancestraljourneys.org/mesolithicdna.shtml|date=2017-04-30}} Mesolithic Western Eurasian DNA</ref>

In a 2015 study published in ''Nature'', the remains of six individuals from Motala ascribed to the Kongemose culture were successfully analyzed. With regards to Y-DNA, two individuals were ascribed to haplogroup I2a1b, one individual was ascribed to haplogroup I2a1, and one individual was ascribed to haplogroup I2c.<ref>{{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 }}</ref>

==Subclades==

The vast majority of individuals belonging to I2 also belong to I-L460, formerly known as I2a. The following subclades are all nested within I-L460.

===I-P37.2=== The '''I-P37.2+''' (also known as '''I2a1a''') (ISOGG 2019). The subclade divergence for I-P37.2 occurred 10.7±4.8 kya. The age of YSTR variation for the P37.2 subclade is 8.0±4.0 kya.<ref name=pmid15162323/> It is the predominant version of I2 in Eastern Europe.<ref>{{cite book |doi=10.1007/978-94-007-6704-1_31 |chapter=The Mediterranean Human Population: An Anthropological Genetics Perspective |title=The Mediterranean Sea |year=2014 | vauthors = Sazzini M, Sarno S, Luiselli D |pages=529–551 |isbn=978-94-007-6703-4 }}</ref> The I2a is further made up by subgroups I-M26, I-M423, I-L1286, I-L880.

====I-L158==== Haplogroup '''I-M26''' (or '''M26''') ''I2a1a1a'' (ISOGG 2019).

Haplogroup I-L158 (L158, L159.1/S169.1, M26) accounts for approximately 40% of all patrilines among Sardinians.<ref>{{cite journal | vauthors = Rootsi S |title=Y-Chromosome haplogroup I prehistoric gene flow in Europe |journal=Documenta Praehistorica |date=31 December 2006 |volume=33 |pages=17–20 |doi=10.4312/dp.33.3 |doi-access=free }}</ref><ref>{{cite journal | vauthors = Francalacci P, Morelli L, Angius A, Berutti R, Reinier F, Atzeni R, Pilu R, Busonero F, Maschio A, Zara I, Sanna D, Useli A, Urru MF, Marcelli M, Cusano R, Oppo M, Zoledziewska M, Pitzalis M, Deidda F, Porcu E, Poddie F, Kang HM, Lyons R, Tarrier B, Gresham JB, Li B, Tofanelli S, Alonso S, Dei M, Lai S, Mulas A, Whalen MB, Uzzau S, Jones C, Schlessinger D, Abecasis GR, Sanna S, Sidore C, Cucca F | display-authors = 6 | title = Low-pass DNA sequencing of 1200 Sardinians reconstructs European Y-chromosome phylogeny | journal = Science | volume = 341 | issue = 6145 | pages = 565–569 | date = August 2013 | pmid = 23908240 | pmc = 5500864 | doi = 10.1126/science.1237947 | bibcode = 2013Sci...341..565F }}</ref> It is also found at low to moderate frequency among populations of the Pyrenees (9.5% in Bortzerriak, Navarra; 9.7% in Chazetania, Aragon; 8% in Val d'Aran, Catalunya; 2.9% in Alt Urgell, Catalunya; and 8.1% in Baixa Cerdanya, Catalunya) and Iberia, and it has been found in 1.6% of a sample of Albanians living in the Republic of North Macedonia<ref>{{cite journal | vauthors = Battaglia V, Fornarino S, Al-Zahery N, Olivieri A, Pala M, Myres NM, King RJ, Rootsi S, Marjanovic D, Primorac D, Hadziselimovic R, Vidovic S, Drobnic K, Durmishi N, Torroni A, Santachiara-Benerecetti AS, Underhill PA, Semino O | display-authors = 6 | title = Y-chromosomal evidence of the cultural diffusion of agriculture in Southeast Europe | journal = European Journal of Human Genetics | volume = 17 | issue = 6 | pages = 820–830 | date = June 2009 | pmid = 19107149 | pmc = 2947100 | doi = 10.1038/ejhg.2008.249 }}</ref> and 1.2% (3/257) of a sample of Czechs.<ref name = "Luca2007">{{cite journal | vauthors = Luca F, Di Giacomo F, Benincasa T, Popa LO, Banyko J, Kracmarova A, Malaspina P, Novelletto A, Brdicka R | display-authors = 6 | title = Y-chromosomal variation in the Czech Republic | journal = American Journal of Physical Anthropology | volume = 132 | issue = 1 | pages = 132–139 | date = January 2007 | pmid = 17078035 | doi = 10.1002/ajpa.20500 | bibcode = 2007AJPA..132..132L | hdl-access = free | hdl = 2108/35058 }}</ref> The age of YSTR variation for the M26 subclade has been calculated as 8.0±4.0 kya.<ref name=pmid15162323/>

====I-L178==== I-L178 is very rare, but has been found in two persons from Germany and one from Poland. The age of YSTR variation for the M423 subclade is 8.8±3.6 kya.<ref name="Peter Underhill 2007 pp. 33-42">{{cite book | vauthors = Underhill PA, Myres NM, Rootsi S, Chow CT, Lin AA, Otillar RP, King R, Zhivotovsky LA, Balanovsky O, Pshenichnov A, Ritchie KH | display-authors = 6 |chapter=New phylogenetic relationships for Y-chromosome haplogroup I: Reappraising its Phylogeography and Prehistory |title=Rethinking the Human Evolution | veditors = Mellars P, Boyle K, Bar-Yosef O, Stringer C |year=2007 |pages=33–42 | publisher = McDonald Institute for Archaeological Research |isbn=978-1-902937-46-5 }}</ref>

====I2a-L621==== thumb|right|The approximate frequency and variance distribution of haplogroup I-P37 clusters, ancestral "Dnieper-Carpathian" (DYS448=20) and derived "Balkan" (DYS448=19: represented by a single SNP I-PH908), in Eastern Europe per O.M. Utevska (2017). I2a1a2b-L621 is typical of the Slavic populations, being highest in Southeastern European regions of Bosnia-Herzegovina and South Croatia (>45%),<ref name=pmid15944443/><ref>{{cite journal | vauthors = Mršić G, Gršković B, Vrdoljak A, Popović M, Valpotić I, Anđelinović Š, Stenzl V, Ehler E, Urban L, Lacković G, Underhill P, Primorac D | display-authors = 6 | title = Croatian national reference Y-STR haplotype database | journal = Molecular Biology Reports | volume = 39 | issue = 7 | pages = 7727–7741 | date = July 2012 | pmid = 22391654 | doi = 10.1007/s11033-012-1610-3 | ref = {{harvid|Mršić et al.|2012}} | s2cid = 18011987 }}</ref><ref>{{cite journal | vauthors = Kovacevic L, Tambets K, Ilumäe AM, Kushniarevich A, Yunusbayev B, Solnik A, Bego T, Primorac D, Skaro V, Leskovac A, Jakovski Z, Drobnic K, Tolk HV, Kovacevic S, Rudan P, Metspalu E, Marjanovic D | display-authors = 6 | title = Standing at the gateway to Europe--the genetic structure of Western balkan populations based on autosomal and haploid markers | journal = PLOS ONE | volume = 9 | issue = 8 | article-number = e105090 | year = 2014 | pmid = 25148043 | pmc = 4141785 | doi = 10.1371/journal.pone.0105090 | doi-access = free | bibcode = 2014PLoSO...9j5090K }}</ref> in Croats (37.7-69.8%), Bosniaks (43.53-52.17%), and Serbs (36.6-42%){{em dash}}often called "Dinaric".<ref name="Fóthi">{{cite journal | vauthors = Fóthi E, Gonzalez A, Fehér T, Gugora A, Fóthi Á, Biró O, Keyser C |title=Genetic analysis of male Hungarian Conquerors: European and Asian paternal lineages of the conquering Hungarian tribes |journal=Archaeological and Anthropological Sciences |volume=12 |issue=1 |date=2020 |article-number=31 |doi=10.1007/s12520-019-00996-0|doi-access=free |bibcode=2020ArAnS..12...31F |quote=We looked at 16 loci from 640 I2a-L621 samples in FTDNA's I2a project database and found that 7 individuals were 2 genetic steps away the Karos samples, of whom 1 was a Hungarian from Kunszentmárton, 2 were Ukrainians, 1 was Lithuanian, 1 was Belarusian, 1 was Russian, and 1 was a German from Poland. Based on SNP analysis, the CTS10228 group is 2200 ± 300 years old. The group's demographic expansion may have begun in Southeast Poland around that time, as carriers of the oldest subgroup are found there today. The group cannot solely be tied to the Slavs, because the proto-Slavic period was later, around 300–500 CE ... The SNP-based age of the Eastern European CTS10228 branch is 2200 ± 300 years old. The carriers of the most ancient subgroup live in Southeast Poland, and it is likely that the rapid demographic expansion which brought the marker to other regions in Europe began there. The largest demographic explosion occurred in the Balkans, where the subgroup is dominant in 50.5% of Croatians, 30.1% of Serbs, 31.4% of Montenegrins, and in about 20% of Albanians and Greeks. As a result, this subgroup is often called Dinaric. It is interesting that while it is dominant among modern Balkan peoples, this subgroup has not been present yet during the Roman period, as it is almost absent in Italy as well (see Online Resource 5; ESM_5). ... Their genetic haplogroup, I2a-CTS10228, is widespread among Slavs, but it is only present in 7% of Caucasian peoples, namely among the Karachay ... As such, it appears that the I2a-CTS10228 haplogroup in the paternal lineage of the Karos leaders arises from a specific branch in the Northern Caucasus dating to about 400–500 CE. Its modern descendents live among the Karachay, Hungarians, and various other surrounding nationalities.}}</ref> It has the highest variance and concentration in Eastern Europe (i.e., Ukraine, Southeastern Poland, Belarus).<ref name="Utevska">{{cite thesis |type=PhD |author=O.M. Utevska |date=2017 |title=Генофонд українців за різними системами генетичних маркерів: походження і місце на європейському генетичному просторі |trans-title=The gene pool of Ukrainians revealed by different systems of genetic markers: the origin and statement in Europe |publisher=National Research Center for Radiation Medicine of National Academy of Sciences of Ukraine|url=http://nrcrm.gov.ua/science/councils/dissertation/|language=uk|pages=219–226, 302}}</ref> According to YFull YTree it formed 11,400 YBP and had TMRCA 6,500 YBP, while its main subclades lineage is I-CTS10936 (6,500-5,600 YBP) > I-S19848 (5,600 YBP) > I-CTS4002 (5,600-5,100 YBP) > I-CTS10228 (5,100-3,400 YBP) > I-Y3120 (3,400-2,100 YBP) > I-Y18331 (2,100 YBP) / I-Z17855 (2,100-1650 YBP) / I-Y4460 (2,100 YBP) / I-S17250 (2,100-1,850 YBP) > I-PH908 (1,850-1,700 YBP).<ref name="YFull">{{cite web|url=https://www.yfull.com/tree/I-PH908/|title=I-PH908 YTree v8.06.01|date=27 June 2020|publisher=YFull.com|access-date=17 July 2020}}</ref>

Older research considered that the high frequency of this subclade in the South Slavic-speaking populations to be the result of "pre-Slavic" paleolithic settlement in the region. Peričić ''et al.'' (2005) for instance placed its expansion to have occurred "not earlier than the YD to Holocene transition and not later than the early Neolithic".<ref name=pmid15162323/><ref name=pmid15944443/><ref>{{cite journal | vauthors = Marjanovic D, Fornarino S, Montagna S, Primorac D, Hadziselimovic R, Vidovic S, Pojskic N, Battaglia V, Achilli A, Drobnic K, Andjelinovic S, Torroni A, Santachiara-Benerecetti AS, Semino O | display-authors = 6 | title = The peopling of modern Bosnia-Herzegovina: Y-chromosome haplogroups in the three main ethnic groups | journal = Annals of Human Genetics | volume = 69 | issue = Pt 6 | pages = 757–763 | date = November 2005 | pmid = 16266413 | doi = 10.1111/j.1529-8817.2005.00190.x | s2cid = 36632274 }}</ref><ref>{{cite journal | vauthors = Rębała K, Mikulich AI, Tsybovsky IS, Siváková D, Džupinková Z, Szczerkowska-Dobosz A, Szczerkowska Z | title = Y-STR variation among Slavs: evidence for the Slavic homeland in the middle Dnieper basin | journal = Journal of Human Genetics | volume = 52 | issue = 5 | pages = 406–414 | date = 16 March 2007 | pmid = 17364156 | doi = 10.1007/s10038-007-0125-6 | doi-access = free }}</ref> However, the prehistoric autochthonous origin of the haplogroup I2 in the Balkans is now considered as outdated,{{refn|group=nb|The SNP I-P37 itself formed approximately 21,000 YBP and had TMRCA 18,400 YBP according to YFull YTree,<ref>{{cite web|url=https://www.yfull.com/tree/I-P37/|title=I-P37 YTree v8.06.01|date=27 June 2020|publisher=YFull.com|access-date=17 July 2020}}</ref> being too old and widespread as an SNP for argumentation of ancient autochthony or medieval migration as well the old research used outdated nomenclature. According to "I-P37 (I2a)" project at Family Tree DNA, the divergence at STR marker DYS448 20 > 19 is reported since 2007,<ref>{{cite web|url=https://www.familytreedna.com/groups/i-2a-hap-group/about/results|title=I2a Y-Haplogroup - Results: I2a2a-Dinaric|publisher=Family Tree DNA|access-date=11 November 2018|quote=Ken Nordtvedt has split I2a2-M423-Dinaric into Din-N and Din-S. Din-N is older than Din-S. N=north of the Danube and S=south of the Danube River ... May 8, 2007: Dinaric I1b1 and DYS 448. DYS448 19 for S and 20 for N.}}</ref> while the SNP which defines the STR Dinaric-South cluster, I-PH908, is reported since 2014.<ref>{{cite web|url=https://i2aproject.blogspot.com/2016/08/link-to-i-l621-tree-showing-major-str_22.html|title=Link to I-L621 tree showing major STR clusters (Updated)|date=22 August 2016|author=Bernie Cullen|work=i2aproject.blogspot.com|publisher=Blogger|access-date=3 April 2019}}</ref> The SNP I-PH908 at ISOGG phylogenetic tree is named as I2a1a2b1a1a1c,<ref>{{cite web|url=https://docs.google.com/spreadsheets/d/1WkRe8UxrhMZ-CaYtPET5rIBbzqCNkGER4YJurbG2ESQ/edit#gid=198726360|title=Y-DNA Haplogroup I and its Subclades - 2019-2020|date=1 October 2019|publisher=ISOGG|access-date=17 July 2020}}</ref> while formed and had TMRCA approximately 1,800 YBP according to YFull.<ref name="YFull"/>}} as already Battaglia ''et al.'' (2009) observed highest variance of the haplogroup in Ukraine, and Zupan ''et al.'' (2013) noted that it suggests it arrived with Slavic migration from the homeland which was in present-day Ukraine.<ref>{{cite journal | vauthors = Zupan A, Vrabec K, Glavač D | title = The paternal perspective of the Slovenian population and its relationship with other populations | journal = Annals of Human Biology | volume = 40 | issue = 6 | pages = 515–526 | date = 2013 | pmid = 23879710 | doi = 10.3109/03014460.2013.813584 | ref = {{harvid|Zupan et al.|2013}} | s2cid = 34621779 }}</ref> O.M. Utevska (2017), in her PhD thesis, despite being a part of research team who came to a different conclusion in 2015,<ref>{{Cite journal |last1=Kushniarevich |first1=Alena |last2=Utevska |first2=Olga |last3=Chuhryaeva |first3=Marina |last4=Agdzhoyan |first4=Anastasia |last5=Dibirova |first5=Khadizhat |last6=Uktveryte |first6=Ingrida |last7=Möls |first7=Märt |last8=Mulahasanovic |first8=Lejla |last9=Pshenichnov |first9=Andrey |last10=Frolova |first10=Svetlana |last11=Shanko |first11=Andrey |last12=Metspalu |first12=Ene |last13=Reidla |first13=Maere |last14=Tambets |first14=Kristiina |last15=Tamm |first15=Erika |date=2015-09-02 |editor-last=Calafell |editor-first=Francesc |title=Genetic Heritage of the Balto-Slavic Speaking Populations: A Synthesis of Autosomal, Mitochondrial and Y-Chromosomal Data |journal=PLOS ONE |language=en |volume=10 |issue=9 |article-number=e0135820 |doi=10.1371/journal.pone.0135820 |issn=1932-6203 |pmc=4558026 |pmid=26332464 |bibcode=2015PLoSO..1035820K |doi-access=free }}</ref> proposed that the haplogroup STR haplotypes have the highest diversity in Ukraine, with ancestral STR marker result "DYS448=20" comprising "Dnieper-Carpathian" cluster, while younger derived result "DYS448=19" comprising the "Balkan cluster" which is predominant among the South Slavs.<ref name="Utevska"/> According to her, this "Balkan cluster" also has the highest variance in Ukraine, which indicates that the very high frequency in the Western Balkan is probably because of a founder effect.<ref name="Utevska"/> Utevska calculated that the STR cluster divergence and its secondary expansion from the middle reaches of the Dnieper river or from Eastern Carpathians towards the Balkan peninsula happened approximately 2,860 ± 730 years ago, relating it to the times before Slavs, but much after the decline of the Cucuteni–Trypillia culture.<ref name="Utevska"/> However, STR-based calculations give overestimated dates,<ref>{{cite journal | vauthors = Šarac J, Šarić T, Havaš Auguštin D, Novokmet N, Vekarić N, Mustać M, Grahovac B, Kapović M, Nevajda B, Glasnović A, Missoni S, Rootsi S, Rudan P | display-authors = 6 | title = Genetic heritage of Croatians in the Southeastern European gene pool-Y chromosome analysis of the Croatian continental and Island population | journal = American Journal of Human Biology | volume = 28 | issue = 6 | pages = 837–845 | date = November 2016 | pmid = 27279290 | doi = 10.1002/ajhb.22876 | quote = It is important to stress that the proposed old age of the I2a1b-M423 and R1a1a1b1a*-M558 lineages obtained in previous studies (Battaglia et al., 2009; Peričić et al., 2005; Rootsi et al., 2004; Underhill et al., 2007, 2015) has been based on STR analysis (8 and 10 loci, respectively) and recent studies clearly indicate that the STR-based age calculations tend to yield overestimated dates (Batini et al., 2015; Hallast et al., 2015; Karmin et al., 2015). | s2cid = 25873634 }}</ref><ref>{{cite journal | vauthors = Balanovsky O | title = Toward a consensus on SNP and STR mutation rates on the human Y-chromosome | journal = Human Genetics | volume = 136 | issue = 5 | pages = 575–590 | date = May 2017 | pmid = 28455625 | doi = 10.1007/s00439-017-1805-8 | s2cid = 3714493 | quote = While the reasons for the difference between genealogical and evolutionary Y-STR rates are thus partly understood, it remains unclear which rate to use. Many have applied the evolutionary rate, though quite a few have used the genealogical, or both, rates. Genetic genealogists generally apply the genealogical rate and criticize population-genetic studies for reporting (in their view) three-times overestimated ages ... The age of each haplogroup was also calculated using the STR genealogical rate and the STR evolutionary rate. Confidence intervals for the two STR-based ages (not shown on the plot) do not overlap. For example, the genealogical age of I2a-L621 (2200 ± 500 years) reaches the envelope age (from 2600 to 3100 ages), while the evolutionary age lies far beyond (9900 ± 2700 years). The observed pattern (Fig. 2a) clearly differs for haplogroups of different age classes. For ages less than 7000 years, the genealogical STR rate provides results consistent with or slightly underestimating the "true" ages, while the evolutionary rate results in three-fold overestimates. For ages between roughly 7000 and 15,000 years neither STR rate provides correct results. For haplogroups older than 15,000 years, the evolutionary rate estimates correctly or overestimates the "true" age. }}</ref> and more specifically, the "Balkan cluster" is represented by a single SNP, I-PH908, known as I2a1a2b1a1a1c in ISOGG phylogenetic tree (2019), and according to YFull YTree it formed and had TMRCA approximately 1,850-1,700 YBP (2nd-3rd century AD).<ref name="YFull"/>

It is considered that I-L621 could have been present in the Cucuteni–Trypillia culture,<ref name="Neparaczki">{{cite journal | vauthors = Neparáczki E, Maróti Z, Kalmár T, Maár K, Nagy I, Latinovics D, Kustár Á, Pálfi G, Molnár E, Marcsik A, Balogh C, Lőrinczy G, Gál SS, Tomka P, Kovacsóczy B, Kovács L, Raskó I, Török T | display-authors = 6 | title = Y-chromosome haplogroups from Hun, Avar and conquering Hungarian period nomadic people of the Carpathian Basin | journal = Scientific Reports | volume = 9 | issue = 1 | article-number = 16569 | date = November 2019 | pmid = 31719606 | pmc = 6851379 | doi = 10.1038/s41598-019-53105-5 | publisher = Nature Research | quote = Hg I2a1a2b-L621 was present in 5 Conqueror samples, and a 6th sample form Magyarhomorog (MH/9) most likely also belongs here, as MH/9 is a likely kin of MH/16 (see below). This Hg of European origin is most prominent in the Balkans and Eastern Europe, especially among Slavic speaking groups. It might have been a major lineage of the Cucuteni-Trypillian culture and it was present in the Baden culture of the Calcholitic Carpathian Basin24 ... The identical I2a1a2b Hg-s of Magyarhomorog individuals appears to be frequent among high-ranking Conquerors, as the most distinguished graves in the Karos2 and 3 cemeteries also belong to this lineage. | bibcode = 2019NatSR...916569N }}</ref> but until now was mainly found G2a and non-I2-L621 clades,<ref>{{cite journal | vauthors = Mathieson I, Alpaslan-Roodenberg S, Posth C, Szécsényi-Nagy A, Rohland N, Mallick S, Olalde I, Broomandkhoshbacht N, Candilio F, Cheronet O, Fernandes D, Ferry M, Gamarra B, Fortes GG, Haak W, Harney E, Jones E, Keating D, Krause-Kyora B, Kucukkalipci I, Michel M, Mittnik A, Nägele K, Novak M, Oppenheimer J, Patterson N, Pfrengle S, Sirak K, Stewardson K, Vai S, Alexandrov S, Alt KW, Andreescu R, Antonović D, Ash A, Atanassova N, Bacvarov K, Gusztáv MB, Bocherens H, Bolus M, Boroneanţ A, Boyadzhiev Y, Budnik A, Burmaz J, Chohadzhiev S, Conard NJ, Cottiaux R, Čuka M, Cupillard C, Drucker DG, Elenski N, Francken M, Galabova B, Ganetsovski G, Gély B, Hajdu T, Handzhyiska V, Harvati K, Higham T, Iliev S, Janković I, Karavanić I, Kennett DJ, Komšo D, Kozak A, Labuda D, Lari M, Lazar C, Leppek M, Leshtakov K, Vetro DL, Los D, Lozanov I, Malina M, Martini F, McSweeney K, Meller H, Menđušić M, Mirea P, Moiseyev V, Petrova V, Price TD, Simalcsik A, Sineo L, Šlaus M, Slavchev V, Stanev P, Starović A, Szeniczey T, Talamo S, Teschler-Nicola M, Thevenet C, Valchev I, Valentin F, Vasilyev S, Veljanovska F, Venelinova S, Veselovskaya E, Viola B, Virag C, Zaninović J, Zäuner S, Stockhammer PW, Catalano G, Krauß R, Caramelli D, Zariņa G, Gaydarska B, Lillie M, Nikitin AG, Potekhina I, Papathanasiou A, Borić D, Bonsall C, Krause J, Pinhasi R, Reich D | display-authors = 6 | title = The genomic history of southeastern Europe | journal = Nature | volume = 555 | issue = 7695 | pages = 197–203 | date = March 2018 | pmid = 29466330 | pmc = 6091220 | doi = 10.1038/nature25778 | bibcode = 2018Natur.555..197M }}</ref><ref>{{cite journal | vauthors = Gelabert P, Schmidt RW, Fernandes DM, Karsten JK, Harper TK, Madden GD, Ledogar SH, Sokhatsky M, Oota H, Kennett DJ, Pinhasi R | display-authors = 6 | title = Genomes from Verteba cave suggest diversity within the Trypillians in Ukraine | journal = Scientific Reports | volume = 12 | issue = 1 | article-number = 7242 | date = May 2022 | pmid = 35508651 | doi = 10.1038/s41598-022-11117-8 | publisher = Nature Research | pmc = 9068698 | bibcode = 2022NatSR..12.7242G }}</ref> and another clade I2a1a1-CTS595 was present in the Baden culture of the Chalcolithic Carpathian Basin.<ref name="Neparaczki"/><ref>{{cite journal | vauthors = Lipson M, Szécsényi-Nagy A, Mallick S, Pósa A, Stégmár B, Keerl V, Rohland N, Stewardson K, Ferry M, Michel M, Oppenheimer J, Broomandkhoshbacht N, Harney E, Nordenfelt S, Llamas B, Gusztáv Mende B, Köhler K, Oross K, Bondár M, Marton T, Osztás A, Jakucs J, Paluch T, Horváth F, Csengeri P, Koós J, Sebők K, Anders A, Raczky P, Regenye J, Barna JP, Fábián S, Serlegi G, Toldi Z, Gyöngyvér Nagy E, Dani J, Molnár E, Pálfi G, Márk L, Melegh B, Bánfai Z, Domboróczki L, Fernández-Eraso J, Antonio Mujika-Alustiza J, Alonso Fernández C, Jiménez Echevarría J, Bollongino R, Orschiedt J, Schierhold K, Meller H, Cooper A, Burger J, Bánffy E, Alt KW, Lalueza-Fox C, Haak W, Reich D | display-authors = 6 | title = Parallel palaeogenomic transects reveal complex genetic history of early European farmers | journal = Nature | volume = 551 | issue = 7680 | pages = 368–372 | date = November 2017 | pmid = 29144465 | pmc = 5973800 | doi = 10.1038/nature24476 | bibcode = 2017Natur.551..368L }}</ref><ref>{{cite journal | vauthors = Patterson N, Isakov M, Booth T, Büster L, Fischer CE, Olalde I, Ringbauer H, Akbari A, Cheronet O, Bleasdale M, Adamski N, Altena E, Bernardos R, Brace S, Broomandkhoshbacht N, Callan K, Candilio F, Culleton B, Curtis E, Demetz L, Carlson KS, Edwards CJ, Fernandes DM, Foody MG, Freilich S, Goodchild H, Kearns A, Lawson AM, Lazaridis I, Mah M, Mallick S, Mandl K, Micco A, Michel M, Morante GB, Oppenheimer J, Özdoğan KT, Qiu L, Schattke C, Stewardson K, Workman JN, Zalzala F, Zhang Z, Agustí B, Allen T, Almássy K, Amkreutz L, Ash A, Baillif-Ducros C, Barclay A, Bartosiewicz L, Baxter K, Bernert Z, Blažek J, Bodružić M, Boissinot P, Bonsall C, Bradley P, Brittain M, Brookes A, Brown F, Brown L, Brunning R, Budd C, Burmaz J, Canet S, Carnicero-Cáceres S, Čaušević-Bully M, Chamberlain A, Chauvin S, Clough S, Čondić N, Coppa A, Craig O, Črešnar M, Cummings V, Czifra S, Danielisová A, Daniels R, Davies A, de Jersey P, Deacon J, Deminger C, Ditchfield PW, Dizdar M, Dobeš M, Dobisíková M, Domboróczki L, Drinkall G, Đukić A, Ernée M, Evans C, Evans J, Fernández-Götz M, Filipović S, Fitzpatrick A, Fokkens H, Fowler C, Fox A, Gallina Z, Gamble M, González Morales MR, González-Rabanal B, Green A, Gyenesei K, Habermehl D, Hajdu T, Hamilton D, Harris J, Hayden C, Hendriks J, Hernu B, Hey G, Horňák M, Ilon G, Istvánovits E, Jones AM, Kavur MB, Kazek K, Kenyon RA, Khreisheh A, Kiss V, Kleijne J, Knight M, Kootker LM, Kovács PF, Kozubová A, Kulcsár G, Kulcsár V, Le Pennec C, Legge M, Leivers M, Loe L, López-Costas O, Lord T, Los D, Lyall J, Marín-Arroyo AB, Mason P, Matošević D, Maxted A, McIntyre L, McKinley J, McSweeney K, Meijlink B, Mende BG, Menđušić M, Metlička M, Meyer S, Mihovilić K, Milasinovic L, Minnitt S, Moore J, Morley G, Mullan G, Musilová M, Neil B, Nicholls R, Novak M, Pala M, Papworth M, Paresys C, Patten R, Perkić D, Pesti K, Petit A, Petriščáková K, Pichon C, Pickard C, Pilling Z, Price TD, Radović S, Redfern R, Resutík B, Rhodes DT, Richards MB, Roberts A, Roefstra J, Sankot P, Šefčáková A, Sheridan A, Skae S, Šmolíková M, Somogyi K, Somogyvári Á, Stephens M, Szabó G, Szécsényi-Nagy A, Szeniczey T, Tabor J, Tankó K, Maria CT, Terry R, Teržan B, Teschler-Nicola M, Torres-Martínez JF, Trapp J, Turle R, Ujvári F, van der Heiden M, Veleminsky P, Veselka B, Vytlačil Z, Waddington C, Ware P, Wilkinson P, Wilson L, Wiseman R, Young E, Zaninović J, Žitňan A, Lalueza-Fox C, de Knijff P, Barnes I, Halkon P, Thomas MG, Kennett DJ, Cunliffe B, Lillie M, Rohland N, Pinhasi R, Armit I, Reich D | display-authors = 6 | title = Large-scale migration into Britain during the Middle to Late Bronze Age | journal = Nature | volume = 601 | issue = 7894 | pages = 588–594 | date = January 2022 | pmid = 34937049 | doi = 10.1038/s41586-021-04287-4 | pmc = 8889665 | bibcode = 2022Natur.601..588P | s2cid = 245509501 }}</ref> Although it is dominant among the modern Slavic peoples on the territory of the former Balkan provinces of the Roman Empire, until now it was not found among the samples from the Roman period and is almost absent in contemporary population of Italy.<ref name="Fóthi"/> According to Pamjav ''et al.'' (2019) and Fóthi ''et al.'' (2020), the distribution of ancestral subclades like of I-CTS10228 among contemporary carriers indicates a rapid expansion from Southeastern Poland, is mainly related to the Slavs and their medieval migration, and the "largest demographic explosion occurred in the Balkans".<ref name="Fóthi"/><ref name="HorolmaTibor2019">{{cite book| vauthors = Pamjav H, Fehér T, Németh E, Koppány Csáji L |title=Genetika és őstörténet|url=https://books.google.com/books?id=xq2xDwAAQBAJ|year=2019|publisher=Napkút Kiadó|language=Hungarian|isbn=978-963-263-855-3|page=58|quote=Az I2-CTS10228 (köznevén "dinári-kárpáti") alcsoport legkorábbi közös őse 2200 évvel ezelőttre tehető, így esetében nem arról van szó, hogy a mezolit népesség Kelet-Európában ilyen mértékben fennmaradt volna, hanem arról, hogy egy, a mezolit csoportoktól származó szűk család az európai vaskorban sikeresen integrálódott egy olyan társadalomba, amely hamarosan erőteljes demográfiai expanzióba kezdett. Ez is mutatja, hogy nem feltétlenül népek, mintsem családok sikerével, nemzetségek elterjedésével is számolnunk kell, és ezt a jelenlegi etnikai identitással összefüggésbe hozni lehetetlen. A csoport elterjedése alapján valószínűsíthető, hogy a szláv népek migrációjában vett részt, így válva az R1a-t követően a második legdominánsabb csoporttá a mai Kelet-Európában. Nyugat-Európából viszont teljes mértékben hiányzik, kivéve a kora középkorban szláv nyelvet beszélő keletnémet területeket.}}</ref> According to a 2023 archaeogenetic study, I2a-L621 is absent in the antiquity and appears only since the Early Middle Ages "always associated with Eastern European related ancestry in the autosomal genome, which supports that these lineages were introduced in the Balkans by Eastern European migrants during the Early Medieval period."<ref name="Olalde2023">{{cite journal |last1=Olalde |first1=Iñigo |last2=Carrión |first2=Pablo |date=December 7, 2023 |title=A genetic history of the Balkans from Roman frontier to Slavic migrations |journal=Cell |volume=186 |issue=25 |pages=P5472–5485.E9 |doi=10.1016/j.cell.2023.10.018 |doi-access=free |pmid=38065079 |pmc=10752003 }}</ref>

Some of the earliest archeogenetic samples until now is Sungir 6 (~900 YBP) near Vladimir, Russia which belonged to the I-S17250 > I-Y5596 > I-Z16971 > I-Y5595 > I-A16681 subclade,<ref name=pmid28982795>{{cite journal | vauthors = Sikora M, Seguin-Orlando A, Sousa VC, Albrechtsen A, Korneliussen T, Ko A, Rasmussen S, Dupanloup I, Nigst PR, Bosch MD, Renaud G, Allentoft ME, Margaryan A, Vasilyev SV, Veselovskaya EV, Borutskaya SB, Deviese T, Comeskey D, Higham T, Manica A, Foley R, Meltzer DJ, Nielsen R, Excoffier L, Mirazon Lahr M, Orlando L, Willerslev E | display-authors = 6 | title = Ancient genomes show social and reproductive behavior of early Upper Paleolithic foragers | journal = Science | volume = 358 | issue = 6363 | pages = 659–662 | date = November 2017 | pmid = 28982795 | doi = 10.1126/science.aao1807 | doi-access = free | bibcode = 2017Sci...358..659S }}</ref><ref>{{cite web|url=https://www.yfull.com/tree/I-A16681/|title=I-A16681 YTree v8.06.01|date=27 June 2020|publisher=YFull.com|access-date=17 July 2020}}</ref> as well I-CTS10228 and I-Y3120 subclades found in two Vikings from Sweden (VK53) and Ukraine (VK542) with predominantly Slavic ancestry of which the second belongs to Gleb Svyatoslavich (11th century).<ref>{{cite journal | vauthors = Margaryan A, Lawson DJ, Sikora M, Racimo F, Rasmussen S, Moltke I, Cassidy LM, Jørsboe E, Ingason A, Pedersen MW, Korneliussen T, Wilhelmson H, Buś MM, de Barros Damgaard P, Martiniano R, Renaud G, Bhérer C, Moreno-Mayar JV, Fotakis AK, Allen M, Allmäe R, Molak M, Cappellini E, Scorrano G, McColl H, Buzhilova A, Fox A, Albrechtsen A, Schütz B, Skar B, Arcini C, Falys C, Jonson CH, Błaszczyk D, Pezhemsky D, Turner-Walker G, Gestsdóttir H, Lundstrøm I, Gustin I, Mainland I, Potekhina I, Muntoni IM, Cheng J, Stenderup J, Ma J, Gibson J, Peets J, Gustafsson J, Iversen KH, Simpson L, Strand L, Loe L, Sikora M, Florek M, Vretemark M, Redknap M, Bajka M, Pushkina T, Søvsø M, Grigoreva N, Christensen T, Kastholm O, Uldum O, Favia P, Holck P, Sten S, Arge SV, Ellingvåg S, Moiseyev V, Bogdanowicz W, Magnusson Y, Orlando L, Pentz P, Jessen MD, Pedersen A, Collard M, Bradley DG, Jørkov ML, Arneborg J, Lynnerup N, Price N, Gilbert MT, Allentoft ME, Bill J, Sindbæk SM, Hedeager L, Kristiansen K, Nielsen R, Werge T, Willerslev E | display-authors = 6 | title = Population genomics of the Viking world | journal = Nature | volume = 585 | issue = 7825 | pages = 390–396 | date = September 2020 | pmid = 32939067 | doi = 10.1038/s41586-020-2688-8 | bibcode = 2020Natur.585..390M | hdl-access = free | hdl = 10852/83989 | biorxiv = 10.1101/703405 | s2cid = 221769227 }}</ref> It was also found in the skeletal remains of Hungarian conquerors of the Carpathian Basin from the 9th century, part of Western Eurasian-Slavic component of the Hungarians.<ref name="Fóthi"/><ref name="Neparaczki"/><ref>{{cite journal | vauthors = Maróti Z, Neparáczki E, Schütz O, Maár K, Varga GI, Kovács B, Kalmár T, Nyerki E, Nagy I, Latinovics D, Tihanyi B, Marcsik A, Pálfi G, Bernert Z, Gallina Z, Horváth C, Varga S, Költő L, Raskó I, Nagy PL, Balogh C, Zink A, Maixner F, Götherström A, George R, Szalontai C, Szenthe G, Gáll E, Kiss AP, Gulyás B, Kovacsóczy BN, Gál SS, Tomka P, Török T | display-authors = 6 | title = The genetic origin of Huns, Avars, and conquering Hungarians | language = English | journal = Current Biology | volume = 32 | issue = 13 | pages = 2858–2870.e7 | date = July 2022 | pmid = 35617951 | doi = 10.1016/j.cub.2022.04.093 | quote = It is notable that the European Y-Hg I2a1a2b1a1a was also specific for the Conqueror group, especially for the elite as also shown before,34 very often accompanied by Asian maternal lineages, indicating that I2a1a2b1a1a could be more typical for the immigrants than to the local population. | s2cid = 249050620 | doi-access = free | bibcode = 2022CBio...32E2858M }}</ref>

===I-M223=== Haplogroup I-M223 aka I2a1b1 (ISOGG 2019), formerly I2a2a (ISOGG 2014). The age of YSTR variation for the I-M223 subclade has been variously estimated as 13.2±2.7 kya,<ref name=pmid15162323/> 12.3±3.1 kya.,<ref name="Peter Underhill 2007 pp. 33-42"/> 14.6 kya<ref>{{Cite web | url=https://yfull.com/tree/I-M223/ | title=I-M223 YTree}}</ref> and 14.6±3.8 kya (Rootsi 2004). I-M223 has a peak in Germany and another in the northeast of Sweden, but also appears in Romania/Moldova, Russia, Greece, Italy and around the Black Sea.<ref>{{cite journal | vauthors = Chiaroni J, Underhill PA, Cavalli-Sforza LL | title = Y chromosome diversity, human expansion, drift, and cultural evolution | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 106 | issue = 48 | pages = 20174–20179 | date = December 2009 | pmid = 19920170 | pmc = 2787129 | doi = 10.1073/pnas.0910803106 | doi-access = free | bibcode = 2009PNAS..10620174C }}</ref> Haplogroup I-M223 has been found in over 4% of the population only in Germany, the Netherlands, Belgium, Denmark, Scotland, and England – also the southern tips of Sweden and Norway in Northwest Europe; the provinces of Normandy, Maine, Anjou, and Perche in northwestern France; the province of Provence in southeastern France; the regions of Tuscany, Umbria, and Latium in Italy; Moldavia and the area around Russia's Ryazan Oblast and Mordovia in Eastern Europe. Of historical note, both haplogroups I-M253 and I-M223 appear at a low frequency in the historical regions of Bithynia and Galatia in Turkey. Haplogroup I-M223 also occurs among approximately 1% of Sardinians.

====I-M284==== Haplogroup I2a1b1a1a (ISOGG 2019) or I-M284, has been found almost exclusively amongst the populations of the United Kingdom and Ireland suggesting that it may have arisen amongst the Ancient Britons, with a most recent common ancestor (MRCA) who lived about 3,100 years BP.<ref>[https://www.yfull.com/tree/I-M284/ YFull, 2021, ''I-M284'']</ref> The presence of this subclade "provides some tentative evidence of ancient flow with eastern areas that could support the idea that the [late Celtic] La Tene culture was accompanied by some migration."<ref name=mcevoybradley/>

Where it is found in those of predominately Irish descent, with Gaelic surnames, it may suggest an ancestor who arrived in Ireland during prehistory, from Celtic Britain.<ref name=mcevoybradley>{{cite book | vauthors = McEvoy BP, Bradly DG |chapter=Irish Genetics and Celts |pages=107–120 | veditors = Cunliffe BW, Koch JT |title=Celtic from the West: Alternative Perspectives from Archaeology, Genetics, Language, and Literature |date=2010 |publisher=Oxbow Books |isbn=978-1-84217-410-4 }}</ref> For example, I-M284 includes many males with the surnames McGuinness and McCartan, who have a single, historically recorded male ancestor in the 6th century; thus it is unlikely to be the result of subsequent migration from Britain to Ireland.<ref name=mcevoybradley/> Some subclades of I-M284 that are atypical of Ireland are relatively common in continental Europe,<ref name=mcevoybradley/> which also supports a point of origin east of Ireland.

====I-CTS10057==== This subclade is commonly referred to as "Continentals". The most recent common ancestor is estimated to have been born around 10,700 years ago.<ref>[https://discover.familytreedna.com/y-dna/I-CTS10057/story FamilyTreeDNA, 2026, ''I-CTS10057'']</ref> It is in turn divided into haplogroups I-L701 (Continental clade 3) and I-Z161 (Continental clades 1 and 2). Continental 3 has a wide distribution. Found in Central Europe from Germany, Austria to Poland, Romania and Ukraine, but also in lower frequencies in Greece, Italy, France, Spain, England, Ireland, and Armenia. It may have been disseminated in part by the Goths. It is nearly absent from Scandinavia and Scotland. The age of I-Z161 is estimated to be around 7,000 years old. It is mainly found in North Europe, especially in Denmark, Germany, the Netherlands, and England. It can also be found in Northwest Sicily.

===I-M436=== {{Expand section|date=January 2021}}

==Structure== Up-to-date phylogenetic trees listing subclades of I2 can be found at YFull.com and FamilyTreeDNA.

== Notes == {{reflist|group="nb"}}

== References == {{Reflist}}

== External links == *[https://www.yfull.com/tree/I2/ YFull YTree of I2] *[https://www.familytreedna.com/public/y-dna-haplotree/I FTDNA Y-DNA Haplotree of I] *[https://docs.google.com/spreadsheets/d/1TH2aUkqHUV8coChJOCxGeXMg6QdyRTPthWOpW8IyCJE/edit#gid=198726360 ISOGG 2019-2020 Y-DNA Haplogroup I and its Subclades]

{{DEFAULTSORT:Haplogroup I2 (Y-Dna)}} I-M438 Category:Human migration Category:Human population genetics Category:Genetic polymorphisms