I-L621 (Y-DNA)

I2a1a2b-L621 is typical of the Slavic populations, being highest in Southeastern European regions of Bosnia-Herzegovina and South Croatia (>45%),[3][13][14] in Bosniaks (43.53-52.17%), Croats (37.7-69.8%), and Serbs (36.6-42%), because of which is often called "Dinaric".[15] It has the highest variance and also high concentration in Eastern Europe (Ukraine, Southeastern Poland, Belarus).[16] 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).[17]

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 place its expansion to have occurred "not earlier than the YD to Holocene transition and not later than the early Neolithic".[2][3][18][19] However the prehistoric autochthonous origin of the haplogroup I2 in the Balkans is now considered as out of date,[nb 1] as already Battaglia et al.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.[24]

The research by O.M. Utevska (2017) concluded 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.[16] This "Balkan cluster" also has the highest variance in Ukraine, which indicates that the very high frequency in the Western Balkan is because of a founder effect.[16] 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.[16] 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).[17]

It is considered that I-L621 could have been present in the Cucuteni–Trypillia culture,[25] but until now was only found G2a,[26] and another subclade I2a1a1-CTS595 was present in the Baden culture of the Chalcolithic Carpathian Basin.[25][27] 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.[15] It was found in the skeletal remains with artifacts, indicating leaders, of Hungarian conquerors of the Carpathian Basin from the 9th century, part of Western Eurasian-Slavic component of the Hungarians.[15][25] According to Horolma 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".[15][28] The earliest archeogenetic sample 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,[29][30] as well I-CTS10228 and I-Y3120 subclades were found in two Viking samples from Sweden (VK53) and Ukraine (VK542) with predominantly Slavic ancestry of which the second belongs to Gleb Svyatoslavich (11th century).[31]

References:

2. (2004). "Phylogeography of Y-Chromosome Haplogroup I Reveals Distinct Domains of Prehistoric Gene Flow in Europe". American Journal of Human Genetics. 75 (1): 128–137.
3. (2005). "High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations". Molecular Biology and Evolution. 22 (10): 1964–1975.
13. (2012). "Croatian national reference Y-STR haplotype database". Molecular Biology Reports. 39 (7): 7727–41.
14. (2014). "Standing at the Gateway to Europe - The Genetic Structure of Western Balkan Populations Based on Autosomal and Haploid Markers". PLOS One. 9 (8): e105090.
15. (2020), "Genetic analysis of male Hungarian Conquerors: European and Asian paternal lineages of the conquering Hungarian tribes", Archaeological and Anthropological Sciences, 12 (1)
16. (2017). Генофонд українців за різними системами генетичних маркерів: походження і місце на європейському генетичному просторі [The gene pool of Ukrainians revealed by different systems of genetic markers: the origin and statement in Europe] (PhD) (in Ukrainian). National Research Center for Radiation Medicine of National Academy of Sciences of Ukraine. pp. 219–226, 302.
17. "I-PH908 YTree v8.06.01". YFull.com. 27 June 2020. Retrieved 17 July 2020.
18. (November 2005). "The Peopling of Modern Bosnia-Herzegovina: Y-chromosome Haplogroups in the Three Main Ethnic Groups". Annals of Human Genetics. 69 (6): 757–763.
19. (16 March 2007). "Y-STR variation among Slavs: evidence for the Slavic homeland in the middle Dnieper basin". Journal of Human Genetics. 52 (5): 406–414.
20. "I-P37 YTree v8.06.01". YFull.com. 27 June 2020. Retrieved 17 July 2020.
21. "I2a Y-Haplogroup - Results: I2a2a-Dinaric". Family Tree DNA. Retrieved 11 November 2018. 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.
22. (22 August 2016). "Link to I-L621 tree showing major STR clusters (Updated)". i2aproject.blogspot.com. Blogger. Retrieved 3 April 2019.
23. "Y-DNA Haplogroup I and its Subclades - 2019-2020". ISOGG. 1 October 2019. Retrieved 17 July 2020.
24. (2013). "The paternal perspective of the Slovenian population and its relationship with other populations". Annals of Human Biology. 40 (6): 515–526.
25. (2019). "Y-chromosome haplogroups from Hun, Avar and conquering Hungarian period nomadic people of the Carpathian Basin". Scientific Reports. Nature Research. 9 (16569): 16569
26. (2018). "The Genomic History of Southeastern Europe". Nature. 555 (7695): 197–203.
27. (2017). "Parallel ancient genomic transects reveal complex population history of early European farmers". Nature. 551 (7680): 368–372.
28. (2019). Genetika és őstörténet (in Hungarian). Napkút Kiadó. p. 58. ISBN 978-963-263-855-3. 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.
29. (2017). "Ancient genomes show social and reproductive behavior of early Upper Paleolithic foragers". Science. 358 (6363): 659–662
30. "I-A16681 YTree v8.06.01". YFull.com. 27 June 2020. Retrieved 17 July 2020.
31. (2019). "Population genomics of the Viking world". Nature. 585 (7825): 390–396

As it is now well known, I2a1 is a typical European haplogroup. It is present all over the continent with maximum frequencies recorded in Bosnia (particularly among Bosnian Croats), Sardinia, Croatia, Serbia (+30%), Montenegro, Romania, Moldova, Bulgaria, and Macedonia (+20%). The percentages of I2a1a2b-L621 clade in Slovenia, Ukraine, and Russia available on the Internet should be taken with caution because they are definitely smaller: up to 13% in Slovenia, 16% in Ukraine, and about 7-8% in Russia. (According to the study of Andrej Zupan /2014/, and data from Ukrainian and Russian DNA projects /2020/) Путь на карте линии I-L621

Haplogroup I2a1a2b-L621, or to be precise, its subclade Y3120, is often considered as another “Slavic” haplogroup (beside R1a-Z282), which is wrong and scientifically inadmissible. The criterion of coherence is unambiguous: “Individuals from genetically distinguishable groups should not be given the same name; individuals from genetically indistinguishable ones should.” 

Haplogroups are a genetic, and not a geographical, linguistic, national, or social category. This means that they cannot be designated according to the language their carriers speak, the country they live in, or religion they practice... Each haplogroup denotes a common genetic origin of the people who share it, and they all belong to the same ethno-racial group which has had its own evolution.

In some cases, haplogroups are easily associated with certain ethno-racial/genetic groups. For example, it is obvious that most of today’s European R1a people (i.e. 99%) are the Slavs, the R1b people are the Celts (or, to be precise: not all R1b men are the Celts, but all Celts are R1b!!), and I1a people are the “Northmen” or Germans in general. Of course, the ancestors of these people were not known by those names 3000 years ago, but they, nevertheless, inherited the same Y-DNA lineages which genetic/ethno-racial Slavs, Celts, and Northmen/Germans inherit today.

Apparently, Y-chromosome studies discovered (and proved) that Slavs and Germans are not just “ethno-linguistic” but also indisputable “ethno-racial” categories – groups of men sharing the same paternal lineages or ancestors. And, for that reason it is necessary to distinguish “I1a Germans” and “R1a Slavs” from the national Germans/Deutsche, Danes, Russians, Poles… (carriers of other haplogroups) who are “Germans” or “Slavs” according to the language they speak (ethno-linguistic) but not according to origin.

Unfortunately, even though the haplogroup I2a is believed to have been the haplogroup of the first anatomically modern humans to inhabit Europe, Cro-Magnons, there is no single common name for all I2a people now. However, "I2a1 people" still represent a separate, distinct ethno-racial group, and they have always been (and will be) "I2a1 people", regardless of the subclade, or nation they belong to, time flow, migrations of their ancestors, place they live in, or the language they speak.

Nations are mixtures of different haplogroups, but haplogroups themselves do not mix. Therefore, not a single I2a1 man participated in the ethnogenesis of the genetic Slavs, but many I2a1 people definitely participated in the ethnogenesis of Slavic (or Germanic, Romance…) speaking nations.

(from http://www.eupedia.com/europe/Haplogroup_I2_Y-DNA.shtml )

According to data collected from an extensive research that has been conducted in the last dozen years, and thousands of ancient and modern DNA test results from different European countries, it seems that the haplogroup I2a-L621 appeared somewhere along the Lower Danube about 12.000 ybp. There is also archeogenetic evidence that some I2a-L621 people lived in the area of today’s Bulgaria 5000 years ago. (ReichLab 2020. SE_Europe; https://haplogroup.info/all-ancient-dna.pdf ) However, the carriers of its oldest lineages have managed to survive until today in central-western Europe. (https://www.yfull.com/tree/I-L621/) No one can tell how or when they arrived in the area, or even if the haplogroup Y3120 (formed 3800 ybp) appeared there or not. At present, it is only confirmed that the “Y3120* men”, negative for downstream branches, live in the Balkans (9 different surnames from Serbia, Bosnia and Herzegovina, Croatia https://dnk.poreklo.rs/DNK-projekat/ ). And, there are some Montenegrins who are yet to be verified.

The initial lack of knowledge and information was confusing and it produced (not less than) two contradicted theories about the place of origin of I-Y3120 and, particularly, its subclades Y4460, Y18331, S17250, and Z17855 which were all formed ~2100 years ago:

a) I2a1 has always been indigenous in the Balkans, and it expanded northward from there;

b) I2a1 originated somewhere in the Poland-Belarus-Ukraine triangle and came to the Balkans with the Slavs in the 6-7th century;

The debate was mostly futile, and many arguments and premises proved to be insignificant, or simply wrong. Namely, the attention was focused on the issues such as haplogroup frequency, the number of representatives, haplotype diversity, archaeological learning of the distant past…, but it turned out that these parameters were unreliable “tools” in locating the area where Y3120 subclades originated, and they were of little help in explaining their development. However, in the last few years, thanks to BigY and SNP test results, the fog has cleared, and it became evident that genetics and history complemented each other.
The first thing that catches the eye of an interested person is the “I2a1 map” provided above. It unmistakably shows that the south-eastern Europe is the “source” of expansion of haplogroup I2a1 (Y3120), but it is not explicit whether it is the eastern or western Balkans. Still, the overall research and some important recent findings go in favour of the eastern side of the peninsula and the Romanian section of the Carpathian Mountains.

Here are some raw facts:

- The carriers of Y3120 subclades live all over south-eastern and north-eastern/north-western Europe. The number of I2a1 people is greater in the north than in the south. But the south-eastern Europe (from Croatia to Greece and Bulgaria) is the only region where all I-Y3120 subclades are clustered together. It is even more significant that the basal lineages of all five Y3120 subclades are found exclusively in the Balkans. (see https://www.yfull.com/tree/I-Y3120/, and Y4460 results at https://bosnjackidnk.com/)The subgroups Y18331, S17250-PH908, and Z17855 are also present in the same area with younger branches and many offsprings – it undoubtedly points to a spatial and temporal continuity of some 2100-2300 years.

Annotation: The larger I2a1 population and the “exclusive” variety of Y4460, S17250-Y4882 and S17250-Y5596 subclades in the “upper” Europe are the consequences of (frequent) migrations over the course of more than 1500 years and, above all, the development of younger branches.

- The newly discovered FGC12098 subclade has only few members so far – from Serbia, Bosnia & Herzegovina, and Poland. (https://www.yfull.com/tree/I-Y3120/ , https://www.familytreedna.com/public/I2aHapGroup/default.aspx?secti...)

- The subclade Y4460 has many representatives in northern Europe, and very few in the Balkans. Some of the Y4460 people in Ukraine, Belarus, Russia… are positive for the initial branches of this subclade, and all Y4460 people in Bulgaria, Macedonia, and Greece belong to a couple of lineages formed 2100 years ago. (The surnames of Y4460 men from western Balkans, who are akin to Bulgarians, can be seen at https://bosnjackidnk.com/baza-bez-markera/)

The diversity of Y4460 in Hungary and Romania is rather conspicuous, and when everything is considered (geographical position of the countries, historical context, the level of presence of this haplogroup in the Balkan countries), it is safe to say that western Dacia was the probable place from which Y4460 spread towards the north, west, and, to a much lesser extent, south.
However, what is even more conspicuous is the following: none of the younger branches (formed less than 2000 years ago), which represent the majority in eastern and western Slavic countries, has been found in the Balkans! This could mean that:

a) Y4460 actually came to eastern Balkan more than 2000 years ago from today’s Slavic countries, remained sparse, and for some reason did not evolve further – which is highly unlikely;

b) the greater part of Y4460 “clan” left the north bank of the Danube in the first centuries CE – they followed the coastline of the Black sea, or the ridges of the Carpathian Mountains and eventually prospered in the Slavic and Baltic countries.

In either case, this is the crucial piece of information: there was no subsequent migration of I-Y4460 people (in the 6th, 7th, or later centuries) to the Balkans which could have brought the younger branches.

- The oldest lineages of Y18331 are found in Greece, Albania, and Macedonia, while others are scattered mainly across the northern and eastern Europe. (https://www.yfull.com/tree/I-Y18331/)It is obvious where this Y3120 branch originated.

- The subclade S17250 was, according to the available data ( id:YF71524SRB, id:YF15878BGR, id:YF14628BIH, id:YF07848BGR, id:YF07821BGR, id:YF06137 HUN) , formed either in the eastern Balkans or somewhere in the Romania-Serbia-Hungary border triangle, but something indeed extraordinary happened 2100-1800 years ago. As it is supposed, this is when the sub-branches PH908, Y4882, and Y5596 appeared. And, for now, it appears that PH908 developed in the western Balkan, and Y4882, Y5596 somewhere over the Danube (from Hungary to Poland).

The following facts indicate that that Y4882 and Y5596 originated in the north: all lineages are there; in the south-eastern Europe Y4882 is represented with only one A1328 branch, and the percentage varies from 0 to 1.5% (of all haplogroups), while Y5596 is practically inexistent. The ancestor(s) of these people could have come to the Balkans with Avars, Slavs..., but also (and more likely) individually, and more recently – because of their small number.

Historically, the appearance of Y4882 and Y5596 can be connected with the Roman expeditions before and at the beginning of the Common Era – some S17250 men from Thrace or Dacia were surely deployed in Germania, but it could also be a result of an independent movement from Pannonia.

It is quite the opposite with PH908 – the basal (12 out of 14) lineages are in the Balkans (https://www. yfull.com/tree/I-PH908/ ); the diversity is remarkable; and it represents the vast majority of I2a1a2b population in Croatia, Bosnia, Serbia, and Montenegro (~2.500.000 men); This is the only Y3120 (S17250) subclade which should be called “Dinaric”.

There are two historical realities which support the Balkan origin hypothesis of PH908.

1) Not many people in Europe know that Montenegro (a small country in the western Balkan) is still a tribal society – people still view themselves as members of different clans which, in most cases, originated from one man. In the late medieval times (13-15th century), Montenegrin tribes were designated as "Arbanases" and "Vlachs" by various (predominantly Serbian) rulers, notaries, and historians. Until the arrival of Turks (the second half of 15th century), Arbanases and, more importantly, Vlachs /Vlahs/ were, without exception, differentiated from Serbs, Croats, or Slavs in general. And, it is established that the medieval Vlachs were descendants of Romanized, pre-Slavic Balkan inhabitants. Recent extensive Y-DNA testing discovered that Arbanas tribes in Montenegro belong to haplogroups E-V13, J2, and R1b, and three major Vlach tribes to I2a S17250-PH908.

2) The presence of PH908 in the central and eastern European countries is mostly the consequence of several well-documented migration waves which began with the Turkish conquest of the Balkans. Namely, it has been established that more than 700.000 people left south-eastern Europe (at least 400.000 of them were from Serbia alone) and settled in Hungarian and Russian empires in the 15th, 17th, and 18th century. Approximately 30-40% of the men belonged to haplogroup I2a1a2b and at least 70% of them carried PH908. This relocation process continued in the 19th century as well and involved thousands of people from Montenegro, Herzegovina, Croatia… It is estimated that the number of their living descendants outside the Balkan Peninsula (Romania excluded) is between 1.500.000 and 2.000.000. (<350.000 in Ukraine, and <650.000 in Russia – calculation performed using the data from Ukrainian and Russian DNA projects /2020/)

- The Z17855 subclade was undoubtedly formed in the eastern Balkans (https://www.yfull.com/tree/I-Z17855/ ) and most of its representatives still live in the same region. It seems that its “mild” expansion (together with the further expansion of I-Y4460, E-V13 and R1b lineages) towards the north started in the 8th century when "the Vlachs crossed the Danube and attacked the residing Slavs". With the arrival of Hungarians these people spread over a wider area. Some Z17855 lineages are confirmed in western Balkans too – for example, the members of the Vlach tribe called Mirilovići from Herzegovina.

_______________________________

The latest ancient Y-DNA study (conducted by Serbian and German scientists), which has not been officially published yet, shows that there is archeogenetic evidence that I2 M423 haplogroup was present in the south-eastern Europe 1500-2000 years ago. Namely, two out of five analysed skeletons from the archaeological site known as Timacum Minus, Kuline necropolis (I-V century CE, Serbia) belonged to that haplogroup. Of course, it is unknown yet if they were L161 or L621.

Be that as it may, the facts stated above indicate that a (small) number of I2a-Y3120 men could have lived in lower Danube region (Hungary, Serbia, Romania) about 2100-2300 years ago. One theory proposes that those people came there from the area in which carriers of primary I2a-L621 lineages live today (Germany, Baden-Württemberg) under the influence of Celts (a tribal confederation called Volcae) who invaded Greece in 279 BCE.

As it happened, one or several of those (Celticized) Y3120 men had (at least) five sons who became the common ancestors of all present-day Y4460, S17250, Z18755, Y18331, and FGC12098 people in the world. In the centuries that followed some of their descendants went to the north-west and some to the south-west. And, that is how modern “Carpathian” and “Dinaric” I2a1a2b branches came into existence.

Scientific studies for additional reading:

http://journals.plos.org/plosone/article?id=10.1371/journal.pone.00... Uniparental Genetic Heritage of Belarusians: Encounter of Rare Middle Eastern Matrilineages with a Central European Mitochondrial DNA Pool

https://www.scribd.com/document/335890846/Genetic-Heritage-of-Croat...

https://medicine.dp.ua/index.php/med/article/view/257 Populations of Transcarpathia and Bukovina on the genetic landscape of surrounding regions (PDF in Russian)

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3590186/ Y-Chromosome Diversity in Modern Bulgarians: New Clues about Their Ancestry

Paleo-Balkan and Slavic Contributions to the Genetic Pool of Moldavians: Insights from the Y Chromosome (Alexander Varzari, 2013)

https://www.biorxiv.org/content/biorxiv/early/2019/04/03/597997.ful... Haplogroups from Hun, Avar and conquering Hungarian period nomadic people of the Carpathian Basin

- This (last) study is rather important because it undoubtedly shows that R1a men and I2a1 men (most probably Y4460 or Z17855) who lived in Panonia (modern Hungary) in the 9th century did not belong to the same ethnic group. Namely, apart from different autosomal and Mt-DNA results, it was discovered that 75%, or 3 out of 4 R1a (Z280-CTS1211) men were lactose tolerant, while 83%, or 5 out of 6/ I2a (xS17250) men were not.