October 11, 2019

I-Y84307 subclade in scientific studies

In the previous article, "PH908 subclades migration", some subclades are based on only two samples, but in the case of the I-Y84307 subclade I-Y93273 both samples are from the same country and as such it cannot be presumed its movement direction to Southeastern Europe. In that regard, this article is an attempt to find the (sub)clade according to the Y-STR haplotype among scientific samples, if possible in populations North of Southeastern Europe.

The scientific studies have a selected list of 12, 17, or 23 markers. However, although due to the presence of Y-STR marker DYS448 (in 17, 23) can be easily predicted I-PH908, the subclades have similar modal haplotypes because of which is difficult to differentiate them without DYS561 and other Y-STR markers. As can be seen in the I-PH908 draft tree (from 23 November 2019), some subclades do have private derivative mutations on Y-STR markers which are characteristic for their prediction, and some of these markers can be found among the selected markers in the studies. Among the selected markers is DYS438 (in 12, 17, 23), which derivation 10 > 9 is seemingly private for I-Y84307 > I-Y93273 (see YFull info).

With that presumption was searched for a PH908 haplotype with an exact mutation on DYS438 which could be a match for the subclade haplotype. It was searched a long list of studies of populations in Central, Eastern, and Southeastern Europe encompassing several thousand samples, including of Albania (Tofanelli 2016), Austria (Berger 2005, Erhart 2012), Belarus (Rębała 2009, Kushniarevich 2013, Pankratov 2016), Bosnia and Herzegovina (Kovačević 2013, Dogan 2016), Bulgaria (Martinez-Cruz 2012, Karachanak 2013), Croatia (Mršić 2012, Šarac 2016, Tofanelli 2016), Czech Republic (Woźniak 2010, Ehler 2011, Purps 2014), Germany (Rodig 2007, Rębała 2013, Purps 2014), Greece (King 2011, Martinez-Cruz 2016, Tofanelli 2016), Hungary (Martinez-Cruz 2012 & 2016, Pamjav 2017), Italy (Purps 2014, Boattini 2015, Tofanelli 2016, Grugni 2018), North Macedonia (Jakovski 2011, Jankova 2019), Moldova (Varzari 2013), Montenegro (Mirabal 2010), Poland (Rębała 2005, Jacewicz 2007-2008, Wolańska-Nowak 2009, Woźniak 2010, Rębała 2013, Purps 2014, Diepenbroek 2019), Romania (Bosch 2006, Stanciu 2010, Bembea 2011, Martinez-Cruz 2012 & 2016), Russia (Roewer 2008, Pesik 2014), Rusyns in Serbia (Rębała 2014, Veselinovic 2014), Serbs in Serbia and near countries (Mirabal 2010, Scorrano 2017, Zgonjanin 2017, Kačar 2019), Slovakia (Petrejčíková 2010, Woźniak 2010, Martinez-Cruz 2016), Slovenia (Zupan 2013, Delser 2018), and Ukraine (Martinez-Cruz 2012 & 2016, Mielnik-Sikorska 2013, Utevska 2015) among others.

A match was found in two samples from Croatia (Mršić 2012), one from Serbia (Mirabal 2010), Hungary (Pamjav 2017), Ukraine (Martinez-Cruz 2012 & 2016), and possibly Romania (Bosch 2006). These six scientific samples were listed with the two samples from YFull currently defining the I-Y84307 > I-Y93273 subclade mainly calculated as one haplotype thus making seven haplotypes in total. In Table 1. are listed haplotypes while in Table 2. are genetic distances between them:

Table 1. List of I-PH908 haplotypes with DYS438=9
Table 2. Genetic distance between the haplotypes

It must be emphasized that Y-STR marker Y-GATA-H4=11 in the AmpFISTR Yfiler PCR Amplification Kit calculation has an equivalent result of Y-GATA-H4=10 in other calculations thus it was edited 11 > 10 in the table because is reducing the initial genetic distance before the tabular inclusion (Table 2.).

Secondly, three haplotypes have or lack some Y-STR markers (Table 1.). While "E818/15" has 23 markers, "UM037" also had DYS426=11 and DYS388=13 which results are shared by YFull haplotype, and "124" also had DYS388=12, DYS436=12, DYS462=12, DYS434=11, DYS461=12, DYS435=11 which results exception of DYS434=9 were shared by YFull haplotype. The latter two additional markers won't be included in the table.

Analysis:

The haplotype of "124" (Romania) has a lack of DYS448=19 to confirm it is I-PH908, but the calculation of Jim Cullen's haplogroup I-subclade predictor gives 40% to be part of Dinaric-South cluster. Additionally, as has a lack of Y-STR markers (13/17) it won't be listed in Table 2., and the specific mutations make its haplotype an isolate, but the closest (4M-5GD) haplotype is of UM037 (Ukraine) with which shares DYS19=17.

According to scientific estimations, the DYS19 has a slow mutation rate, while DYS437 and DYS438 have an even slower rate. This is significant because of the genetic distance between the haplotypes in comparison to "104" (Serbia), ranging from 4M-4GD to 6M-8GD on only 17 Y-STR markers. The sample "104" with DYS437=14, DYS19=16, DYS439=12, DYS458=18, DYS635=23, and even DYS385ab=15-15 does not indicate to be related with other haplotypes, neither as distant branching of the subclade. It is rather a part of a modal haplotype of another currently unknown subclade which is defined by some or most of these marker results (see e.g. SDP). This indicates that the rare STR marker result DYS438=9 is not exclusively present in the I-Y84307 subclade and problematizes the characteristic definition for the same.

As for the genetic distance between the three haplotypes from Croatia, the "71" (Central Croatia) and "72" (Southern Croatia) are very close to the YFull haplotype (1M-2GD, 2M-2GD), and probably could belong to the I-Y84307 subclade. According to YFull and FTDNA analysis of SNPs and STRs, the most recent common ancestor of the two YFull samples is dated to 1250 or 1050, as such the MRCA of these haplotypes lived around or before that time period. The ancestral location was probably located somewhere in a triangle of Western, Central, and Southern regions (Fig. 1). Like the YFull haplotype, the "72" has the same close genetic distance to "E818/15" and "UM037" (2M-2GD, 2M-3GD), while "71" is less close (3M-4GD, 3M-5GD).

Fig. 1 Map of Croatian regions, Mršić et al. 2012

The "E818/15" (Hungary; Bodrogköz, Zemplénagárd) and "UM037" (Ukraine; c. Ivano-Frankivsk and Chernivtsi Oblast) haplotypes are very close to each other (1M-1GD) with an only difference on DYS19. Interestingly enough, the localization of these two haplotypes, with the addition of "124" (Romania; Ploiești), has geographical proximity and gravitation in Eastern Europe (Fig. 2). With the presumption that these three haplotypes are related to the haplotypes from Croatia, the location could suggest the medieval place of origination and movement to the Western Balkans.

Fig. 2 Map of I-Y84307 with the location (stars) of the three DYS438=9 samples

Genealogical database samples:

Besides the initially mentioned YFull samples, in genealogical databases exist three more I-PH908 sample haplotypes with DYS438=9. Their haplotypes won't be displayed but will be compared for genetic distance (Table 3.). First is "T1851" from the Czech Republic with 17 markers (see CDP), the second is "101953" from Montenegro with 23 markers and belongs to the I-Y52621 subclade (see SDP), while third is "142396" from Greece with 67 markers (see GDP). As shown, at least two other subclades have one rare sample with a private mutation on DYS438. That problematizes the evaluation of a relationship between haplotypes and whether they belong to the I-Y84307 subclade. However, presuming the relationship and in the context of being in a population located North of Southeastern Europe, on the map is added: "T1851" (Plzeň) which could suggest a different place of origination and movement (Fig. 3).

Table 3. Genetic distance between the haplotypes
Fig. 3 Map of I-Y84307 with the location (stars) of the four DYS438=9 haplotypes

Y Chromosome Haplotype Reference Database samples:

As most of the scientific haplotypes are in YHRD it will be taken into account 1-2-step neighbor haplotypes of the previously mentioned haplotypes, specifically DYS438=10 with the assumption it is an ancestral mutation of the derivative DYS438=9 and that DYS438=10 haplotypes could also belong to I-Y84307 > I-Y93273 (Table 4.). The haplotypes location, including North of Southeastern Europe, was added on the map (Fig. 4), additionally to them neighbors of "T1851" which is not listed in the haplotype table, while "101953" has closest neighbors from Southeastern Europe. The "101953" and "104" are not placed on the map because of they have private mutations of another subclade. The "142396" lacks the DYS635 marker but nevertheless the result due to private mutations didn't have a match and since it is not known the exact location it was putatively located (blue star). The mentioned "T1851" had 1-step neighbor DYS438=10 from Ukraine (Lviv), Slovenia (Ljubljana) and Croatia (Zagreb), while the 2-step neighbors North of Southeastern Europe in Poland (Warsaw), Germany (Stuttgart) and Russian Federation (Ivanowo).


Table 4. List of I-PH908 1-2-step-neighbor haplotypes with DYS438=9 and DYS438=10
Fig. 4 Map with the location of listed I-PH908 1-2-step-neighbor haplotypes DYS438=9 (star) and DYS438=10 (triangle)

Conclusion:

In conclusion, the STR marker result DYS438=9 probably is indicative for the PH908 subclade I-Y84307 > I-Y93273, but only if are taken into account results of other STR markers which preceded it like DYS561=15, and are present or absent in the modal haplotype because it is a rare private mutation found in other PH908 subclades. Further STR and SNP sampling are needed to definitively confirm whether the specific marker result is characteristic for all the eight SNPs currently defining the subclade. Among many scientific and genetic genealogical samples were found two very close contemporary haplotypes, both in Croatia, indicating that the nearest genetic matches most probably live in the same country. In short, if it is possible depending on the haplotype of any haplogroup, it is advised to anyone to check both commercial & genealogical and scientific databases & studies for a match as it could give further information on a geographical and historical ancestry.

July 1, 2019

PH908 subclades migration

In the previous and multi-disciplinary article, "South Slavic origin and I2a-Dinaric South (PH908)", it was concluded that the main movement of the Slavs to Southeastern Europe was from an area in Eastern Europe, roughly Eastern Carpathians and Western Ukraine. In regard to genetic studies results that were primarily based on the frequency and variance of contemporary populations, this article will have a different approach.

Isofrequency contour map for haplogroup I2-CTS10228, see the full resolution

A map based on archaeological and linguistic data. For a high-quality map see Scribble Maps

Considering the conclusion that the haplogroup I-PH908 arrived in the Early Middle Ages from the North this post will be a synthesis of the countries of origin of the I-PH908 subclades found at the YFull YTree v9.02.00, Y-DNA Haplotree, and Y-haplogroup I2a Project (including a post and post from 26 January 2021) at FTDNA. The information will be used to make a simplified draft tree, approximately visually localize them North of Southeastern Europe, and presume the subclades movement to Southeastern Europe. However, it is emphasized that the information is based on a very limited and biased number of samples who made the NGS test and does not necessarily reflect the factual reality in accordance with scientific frequency and diversity due to sampling bias. If each PH908 individual in Eastern and Central Europe would have been tested, due to the old formation and TMRCA age of subclades and participation in the Early Slavic migrations, probably all of them are more or less scattered in both parts of Europe.

DYS561=15:

0) I-PH908*: Albania, "Czech Republic", "Germany", "Latvia", Poland, Russia, Serbia etc.

1) A356/Z16983*: "Hungary", "Poland", Albania, Serbia, United States
* Y110105*: Germany
** Y97246: Russian Federation/Finland
*** BY188785: Czech Republic, Poland
* A493*: Albania, Bosnia and Herzegovina, "Croatia", "Germany", "Greece", North Macedonia, "Montenegro", "Serbia", Slovakia, "Ukraine", USA
** Y6651/Y6652: Czech Republic
** Y55893: Montenegro
*** Y45843: Bosnia and Herzegovina, Croatia
** BY169115: Italy, Serbia
** FT69508/FT69595: Croatia, Montenegro
** Y111673: Germany, Finland (?), USA
** A8740/A8741*: Germany, Poland, USA
*** Y59865: United States

Z16983
2) A5913/A5914*: Bosnia and Herzegovina, Croatia, Hungary
* A22312*: Montenegro, Poland
** BY153567
*** BY154460/BY152858: Montenegro
*** Y132790: Bosnia and Herzegovina
*** FT65958
**** FT65952: Montenegro, "Yugoslavia"
* A16863: Finland, Russian Federation, Ukraine

A5913
3) Y32084
* Y140541: Greece
** BY227002: Bulgaria
* PH3310/A14301*: Ukraine
** A20333: Kosovo, Ukraine
*** FT14649: Serbia
**** S10860: Bosnia and Herzegovina
** PH1012/PH3314: Serbia

Y32084
4) Y84307:
*BY87256/Y93273*5+SNPs: Croatia
*FTA65128: Serbia, Turkey

5) FT168731/FT169458/FT191198*: Poland
* FT168415*10+SNPs: Tatarstan

FT168731
6) BY198275
* FT162921: Czech Republic, Poland
* P41.2&M359.2/BY200456: Bosnia and Herzegovina // Istanbul, Turkey (Cinnioglu et al. 2004), Bosnia and Croatia (Battaglia et al. 2009), Bulgaria and Romania (Martinez-Cruz et al. 2012)

BY198275
7) FT275645/Y200082*6+SNPs: Croatia, Serbia

8) BY93199: Montenegro

9) FT277965: North Macedonia
* YP206: Sardinia (Francalacci et al. 2013)

10) Y178551: Greece [Macedonian] (DYS561=n/a)

11) Y93865: Bulgaria, Greece (DYS561=n/a)

DYS561=16

0) I-PH908*: Bulgaria, Montenegro, Germany, "Poland", "Ukraine", Serbia etc.

1) FT16449*: "Bosnia and Herzegovina", Bulgaria, Russian Federation
* Y126296: Ukraine/Poland
** FT19919: Bosnia and Herzegovina
** Y81557: Serbia
*** BY57773: Serbia
**** Y189253/FT240062
***** FT354262: Croatia, Italy (Albania)
**** Y103938: Bosnia and Herzegovina
***** Y99196: Croatia
* FT36524: Belarus, Russian Federation
* MF2888: Croatia, Germany, Montenegro
* BY173300
** BY67102: Bosnia and Herzegovina, Croatia
* FT138628: Bulgaria, Serbia
* Y109702: Czech Republic
** Y99608: Germany, Poland
* Y151633
** FT33812: United States (Croatia), Croatia
** BY169079: Bosnia and Herzegovina
*** BY171718: Croatia
**** Y144303
***** Y144305: Austria, Croatia
***** A22770/BY170155: Croatia
* BY97555//Y51673: Bosnia and Herzegovina, Slovakia
* FGC44297/FT166563: Bulgaria
* Y57291/Y46636: Bosnia and Herzegovina, Serbia

FT16449

Y51673

2) FT14506*: Bosnia and Herzegovina, Bulgaria, Hungary, Serbia
A13912: Bulgaria
** A13907/Y30729/BY37511: Croatia, "Hungary"
*** BY106777: Croatia
* FT177529: Poland
** FT176058: Russian Federation
*** FT260408: Poland (Galicia)
**** FT261843: Poland
***** FT259981: Poland
* BY135769/BY14506: Poland (Galicia), Russian Federation
* FT41224: Bulgaria
** Y52621: Bosnia and Herzegovina, Moldova
*** FT190799: Montenegro, Serbia
** BY191170/BY191770: Croatia
*** FT271768: Italy, "Serbia"
*** BY189804: Bulgaria
**** BY189748: "Serbia"
***** BY190900: Bosnia and Herzegovina, Ukraine
* Y56203*: Poland (Galicia)
** FT25907: Bosnia and Herzegovina
** Y134603/Y134578: Montenegro, Serbia
* BY186407: Bulgaria, Montenegro

FT14506

When all the maps are merged it indicates that there's a higher variance of I-PH908 subclades in Eastern Europe than in Central Europe and that from Eastern Europe migrated in the direction of West and South. Most if not all subclades arrived in Southeastern Europe from Eastern Europe, roughly along the area of Eastern Carpathian Mountains and through Carpathian Basin, most probably during the period of Pannonian Avars and revolts against them by Slavs and Bulgars by the mid-7th century. It is not excluded a possibility of a very small Polabian Slavic contribution (only for some subclades of Z16983 and FT16449). The samples belonging to currently unknown subclades are also largely from the countries in Eastern Europe, while in Southeastern Europe particularly Bulgaria, which also indicates the movement's direction. The result overwhelmingly supports the conclusions from the previous post.

A merged map with arrows of migration and subclades list

Presumed movement of the Slavs along the edges of Carpathians, and through Moravian Gate in the West, considering geographical relief but assuming it was only through lowlands and excluding all mountain passes (e.g. Verecke Pass)

Presumed movement of the Slavs along the edges of Carpathians, and through Moravian Gate in the West, considering geographical relief, but assuming it was both through lowlands and mountain passes

"" - Mean the sample is confirmed based on Y-STR and single SNP test results
// - Mean the listed samples after // are confirmed in scientific studies
* The post will be updated every couple of months with the development of the Y-Tree (27.01.2021.).

April 10, 2019

South Slavic origin and I2a-Dinaric South (I-PH908)

In this article are gathered up to date scientific information about the medieval (South) Slavic migration to Southeastern Europe which is dated at least since the mid-6th and early 7th centuries. The information is from various scientific fields, like historiography, archaeology, linguistics, anthropology and population genetics, being a contemporary multi-disciplinary approach and synopsis on such a topic. The population genetics focus will be first on autosomal DNA, to establish an overview of the ethnogenesis of the contemporary populations before and after the medieval migration, and then on Y-DNA and particularly the distribution and possible medieval migration pattern of the haplogroup I2 > I-PH908 in the male population. At the end is a summary of the conclusions. The article was last updated on 25 October 2024.

1. Autosomal DNA:

In this part will be cited and made an analysis of contemporary autosomal DNA results from Ralph et al. (2013), Hellenthal et al. (2014), Kovacevic et al. (2014), Kushniarevich et al. (2015), Delser et al. (2018) among others and share additional independent research of recent archaeogenetic samples.

According to The Geography of Recent Genetic Ancestry across Europe by Ralph et al. 2013;

- the IBD method shows that "Southeastern Europeans, for example, share large numbers of common ancestors that date roughly to the era of the Slavic and Hunnic expansions around 1,500 years ago", specifically, the green bar in the leftmost panels tell us that Serbo-Croatian speakers substantially share common ancestors with the sampled Polish and Romanian-Bulgarian speakers in the medieval period (500-1500 YBP) and the ancient period (1500-2500 YBP).

- the IBD method correlates with "the expansion of the Slavic populations into regions of low population density beginning in the sixth century, reaching their maximum by the 10th century. The eastern populations with high rates of IBD are highly coincident with the modern distribution of Slavic languages, so it is natural to speculate that much of the higher rates were due to this expansion. The inclusion of (non-Slavic speaking) Hungary and Romania in the group of eastern populations sharing high IBD could indicate the effect of other groups (e.g., the Huns) on ancestry in these regions, or because some of the same group of people who elsewhere are known as Slavs adopted different local cultures in those regions. Greece and Albania are also part of this putative signal of expansion, which could be because the Slavs settled in part of these areas (with unknown demographic effect), or because of subsequent population exchange"

Ralph et al. 2013

According to A Genetic Atlas of Human Admixture History by Hellenthal et al. 2014;

- the Globetrotter shows that "the other two source groups appear much more local. One is more North-European in the repainting - when we exclude other East European groups as donors - and is largely replaced by northern Slavic-speaking groups in our original analysis. The other source is more southerly (e.g. Greece, West Asians). This local migration could explain a recent observation of an excess of IBD sharing in Eastern Europe – including in the Greeks, in whom we infer admixture involving a group represented by Poland, at the same time - that was dated to a wide range between 1,000 and 2,000 years ago. We speculate that these events may correspond to the Slavic expansion across this region at a similar time, perhaps related to displacement caused by the Eurasian steppe invaders"

- another analysis "with the main admixture event being the Slavic expansion and hereof, on a consistent to all receipents indication of the Slavic expansion by analyzing only a single Slavic-speaking population (Polish) as either of the two donor populations of the recipient populations. The authors speculate the Polish ancestral component may correspond to the Slavic expansion. The second donor group of the recipient populations is random and not consistent at all" dated to 500-900 CE or a bit later with over 40-50% among Bulgarians, Romanians, and Hungarians.

Hellenthal et al. 2014


- the Admixture analysis of autosomal variation shows that "the most illustrative population structure for the populations of the Western Balkan area is achieved at K = 7, with three dominant ancestral components. Beside the most apparent dark blue European component, a largely South/West-European-specific light blue and a beige component, shared mostly with the populations from the Caucasus and the Middle East are observed" and that "profiles of all three ethnic groups of Bosnia are almost identical"

- the Principal Component Analysis and Fst distances of autosomal variation show that "the resulting network exemplifies genetic affinity between Western Balkan populations that form a bridge between East-European Slavic speakers and populations from Eastern Balkan and the Middle East. The Croatians and Bosnians are more close to East European populations and largely overlap with Hungarians from Central Europe, while Kosovars and Macedonians cluster closer to Eastern Balkan populations and Gagauzes" but the "Kosovars deviate the most from other Western Balkan populations – note, that among those they have also the biggest similarity to Greeks. Serbians and Montenegrins have an intermediate position on PCA plot and on Fst –based network"

- the TreeMix analysis of the same subset of populations used in Admixture analysis shows that "The Western Balkan populations take central position on the tree and are surrounded by the Eastern Balkan and South European populations from one and the Eastern Slavic populations together with Poles and Hungarians from the other side. The latter three form the tipmost branch of the tree. The migration events with the highest weight are directed towards the Eastern Balkan populations – to Romanians (migration weight 0.49) and to Bulgarians (weight 0.47), who have received the considerable gene flow from the root of the edge encompassing East Slavic populations, Poles, Hungarians and Bosnians from Western Balkan"

- the Variation of haploid markers of Western Balkan populations shows that "the Western Balkan populations are closest to their Slavic-speaking neighbours both according to maternal (Czechs and Belarusians) and paternal (Slovaks) variation, but it has to be noted that the pooled sample is biased towards northern populations of Western Balkan (Bosnia and Herzegovina, Croatia) and thus represents mostly the variation of this part of the study region. In autosomal analysis, the Bosnians and Croatians are closest to Hungarians, the East Europeans and Eastern Balkan populations are at the same distance from these Western Balkan populations. East European Slavic-speakers are similar to our pooled Western Balkan sample of PCA also in mtDNA and NRY analyses and the Hungarians in NRY analysis"

Focusing on the first paragraph "Admixture analysis of autosomal SNPs of the Western Balkan region in a global context on the resolution level of 7 assumed ancestral populations", see image below, will be made an approximate numerical percentage with a digital ruler of samples from Hungary (19), Croatia (13), Serbia (14), and Bulgaria (13).

Kovacecic et al. (2014)

In low percentages are:
*Green - Southern Asia and Near East, calculated as "south"
*Yellow - Eastern Asia and somehow Northern Eurasia, calculated as "north" due to the presence among sampled North Russians
*Orange - Near East, calculated as "south"

In median and high percentages are:
*Red - Caucasus and Anatolia, calculated as "south"
*Light Blue - Southern and Western Europe characterized by isolated populations of Sardinia and Basques, calculated as "south"
*Dark Blue - Eastern Europe characterized by Slavic populations, calculated as "north"

Horizontally listed are Hungary, Croatia, Serbia, and Bulgaria:
*Green (H: 1.185%), (C: 1.69%), (S: 0%), (B: 0%)
*Yellow (H: 1.185%), (C: 0%), (S: 0.86%), (B:1.69%)
*Orange (H: 0%), (C: 0%), (S: 0%), (B: 0.84%)
*Red (H: 21.35%), (C: 22.03%), (S: 31.35%), (B: 35.59%)
*Light Blue (H: 15.25%), (C: 15.25%), (S: 16.94%), (B: 16.10%)
*Dark Blue (H: 61.01%), (C: 61.01%), (S: 50.84%), (B: 45.76%)

In total:
*North (H: 62.19%), (C: 61.01%), (S: 51.68%), (B: 47.45%)
*South (H: 37.78%), (C: 38.97%), (S: 48.29%), (B: 52.53%)

The 2014 study indicates that the specific contemporary populations have approximately at least 45% up to 60% shared ancestry with Northern populations predominantly of Slavic origin, roughly said medieval migratory ancestry, and at least 35% up to 50% shared ancestry with Southern populations, roughly said ancient autochthonous ancestry.

According to Genetic Heritage of the Balto-Slavic Speaking Populations: A Synthesis of Autosomal, Mitochondrial and Y-Chromosomal Data by Kushniarevich et al. 2015, see image below;

- the data shows that "Most South Slavs are separated from the rest of the Balto-Slavic populations and form a sparse group of populations with internal differentiation into western (Slovenians, Croatians and Bosnians) and eastern (Macedonians and Bulgarians) regions of the Balkan Peninsula with Serbians placed in-between"

- the Admixture analysis was "modeled ancestral genetic components in Balto-Slavic populations. Assuming six ancestral populations (K = 6), Balto-Slavic speakers bear membership almost exclusively from two ancestral components: the dark blue (k3) and the light blue (k2), albeit in different proportions. k3 is omnipresent throughout European populations and decreases from north-eastern Europeans southwards. Thus, k3 peaks in Baltic speakers and prevails in East Slavs (80–95%) and decreases notably in South Slavs (55–70%). In contrast, k2 is abundant around the Mediterranean and in the Caucasus region and decreases among Europeans when moving northward. Accordingly, it makes up nearly 30% of ancestral proportions in South Slavs, decreases to around 20% in West and East Slavs and drops to around 5% in North Russians and Baltic speakers"

- the analysis of IBD segment distributions shows "South Slavs in their turn share a similar number of IBD segments with East-West Slavs and with the ‘inter-Slavic’ Romanian, Hungarian and Gagauz populations. Notably, South Slavs share significantly fewer IBD segments for length classes 1.5–3 cM with their immediate geographic neighbors in south–Greeks–than with the group of East-West Slavs" but "revealed even patterns of IBD sharing among East-West Slavs–‘inter-Slavic’ populations (Hungarians, Romanians and Gagauz)–and South Slavs, i.e. across an area of assumed historic movements of people including Slavs" with a presence of "influence of geography in shaping the Slavic genetic heritage" and that "taken together, several mechanisms including cultural assimilation of the autochthonous populations by expanding Slavs while maintaining the pre-Slavic genetic boundaries, and in situ gene pool shaping, are needed to explain the genetic patterns observed on the eastern, north-eastern and western margins of the current ‘Slavic area’ within Central-East Europe. The presence of two distinct genetic substrata in the genomes of East-West and South Slavs would imply cultural assimilation of indigenous populations by bearers of Slavic languages as a major mechanism of the spread of Slavic languages to the Balkan Peninsula."

Kushniarevich et al. 2015

According to the Genetic Landscape of Slovenians: Past Admixture and Natural Selection Pattern by Delser et al. 2018;

- the Admixture analysis of "Slovenian samples show an admixture pattern similar to the neighboring populations such as Croatians and Hungarians"

- the Principal Component analysis of "first two PCs explained ∼16% of the variance with Slovenian samples grouping together with the Croatians, Hungarians and close to the Czechs... All Slovenian samples group together with Hungarians, Czechs, and some Croatians (Central-Eastern European cluster)"

- the analysis of the "UPGMA tree based on the Fst matrix shows all Slovenian individuals clustering together with Hungarians, Czechs, Croatians, Ukrainians, and Belarusians... Pattern of runs of homozygosity computed on the Slovenian population does not differ significantly from Hungarians, Czechs, Croatians (Mann Whitney p-value > 0.09), albeit showing a slightly higher amount of segments and total homozygosity"

Focusing on the first paragraph "Unsupervised admixture analysis" on the resolution level of 5 assumed ancestral populations, see the image below, will be made an approximate numerical percentage with a digital ruler of samples from Slovenia, Hungary, Croatia, and Romania.

Delser et al. 2018
In low percentages are:
*Yellow - Southwestern Europe characterized by isolated Basques population, "south"
*Red - Southwestern Europe characterized by isolated Sardinian population, "south"

In median and high percentages are:
*Green - Southeastern Europe, "south"
*Light Blue - Northwestern Europe, "north"
*Dark Blue - Northeastern Europe, characterized by Slavic populations, "north"

Horizontally listed are Hungary, Slovenia, Croatia, and Romania:

*Yellow (H: 5.08%), (S: 5.93%), (C: 3.38%), (R: 3.38%)
*Red (H: 6.77%), (S: 6.77%), (C: 6.77%), (R: 9.32%)
*Green (H: 23.72%), (S: 23.30%), (C: 32.20%), (R: 47.45%)
*Light Blue (H: 23.72%), (S: 22.03%), (C: 20.33%), (R: 11.86%)
*Dark Blue (H: 40.67%), (S: 41.94%), (C: 37.28%), (R: 27.96%)

In total:
*North (H: 64.39%), (S: 63.97%), (C: 57.61%), (R: 39.82%)
*South (H: 35.57%), (S: 36%), (C: 42.35%), (R: 60.15%)

The 2018 study indicates that the specific contemporary populations have approximately at least 40% up to 65% shared ancestry with Northern populations predominantly of Slavic origin, with a south-north gradation, roughly said medieval migratory ancestry, and at least 35% up to 60% shared ancestry with Southern populations, with a north-south gradation, roughly said ancient autochthonous ancestry.

According to the Inferring Recent Demography from Isolation by Distance of Long Shared Sequence Blocks by Ringbauer et al. 2017, was "detected a systematic, large-scale deviation from a simple diffusion model with uniform population density, as there is a clear gradient for higher block sharing in the direction of the Balkan countries (Figure 5). This was already observed by Ralph and Coop (2013). They hypothesized that this could be due to the historic Slavic expansion, a hypothesis supported by admixture analysis (Hellenthal et al. 2014). However, the pattern of increased block sharing also holds for longer, typically younger, blocks, which could hint additionally at a consistently lower population density in these regions. Such systematic regional deviations from the diffusion model also imply that care should be taken when estimating parameters and their uncertainty ranges". According to the Estimating recent migration and population-size surfaces by Al-Asadi et al. 2019, the "inferred population size surfaces for the POPRES data show a general increase in sizes through time, with small fluctuations across geography; In our results, the smallest inferred population sizes are in the Balkans and Eastern Europe more generally. This is in agreement with the signal seen previously [Ralph 2013]; however, taken at face value, our results suggest that high PSC sharing in these regions may be due more to consistently low population densities than to historical expansions (such as the Slavic or Hunnic expansions). Although consistent with previous results, our estimates of dispersal and population sizes do not exactly agree with empirical estimates. For example, our estimates of population sizes are consistently lower than the census sizes (S5 Fig)". According to the Diversity and Structure of Mitochondrial Gene Pools of Slavs in the Ethnogenetic Aspect by Malyarchuk & Derenko 2021, "according to archaeological and paleoanthropological data, the settlement of the Slavs from the ancestral territory, which was presumably located in central Europe, occurred more intensively in the early Middle Ages (in the 6th–7th centuries AD) and was accompanied by genetic interaction with the peoples who inhabited the territories of the migration routes of Slavs ... analysis of large samples of mitogenomes in the populations of East, West, and South Slavs indicated the regional specificity of Ne dynamics: a rapid increase in population size during the Bronze Age in eastern Europe, during Eneolithic Age in central Europe, and during the early Middle Ages in southern Europe". According to the Revealing the recent demographic history of Europe via haplotype sharing in the UK Biobank by Gilbert at al. 2022, "we observe NW Balkans presenting slightly longer within-cluster IBD segments than NE Balkans, which is matched with a consistently lower Ne and elevated ROH—suggestive of a smaller population than the northeast of the Balkans or neighboring central Europe to the north". According to the Reconstructing the history of founder events using genome-wide patterns of allele sharing across individuals by Tournebize et al. 2022, also confirmed historically recent founder effect in the Balkans.


- "Individuals from the Kuline necropolis of Timacum Minus places them in the 10th century CE ... In light of these results, we modeled the ancestry of the Kuline individuals as a mixture of 56% deriving from the local Balkan Iron Age substratum and 44% deriving from Northeastern European Iron Age groups, and obtained a good statistical fit (Figure 2; Supplementary section 12.8). Our results point to a strong demographic impact of Eastern European groups in the Balkans during the Medieval period, likely associated to the arrival of Slavic-speaking populations. Yet, our results rule out a complete demographic replacement, as we observe a significant portion of local Iron Age Balkan ancestry in Kuline individuals."

- "Present-day Serbs, Croats and the rest of central/northern Balkan populations yielded a similar ancestral composition as the Kuline individuals, with approximately 50% Northeastern European-related ancestry admixed with ancestry related to Iron Age native Balkan population (Figure 3), implying substantial population continuity in the region over the last 1,000 years. This ancestry signal significantly decreases in more southern groups, but it is still present in populations from mainland Greece (~30%) and even the Aegean islands (7-20%)."

Olalde et al. 2021: Proportions of Northeastern European-related ancestry (in black) for present-day Balkans populations. 10th-century Kuline individuals are indicated in a red circle

According to the newly published study A genetic probe into the ancient and medieval history of Southern Europe and West Asia by Lazaridis et al. 2022, Slavic migrations "profoundly affected the region";

- "We highlight Roman, Byzantine, and medieval individuals from Albania, Bulgaria, Croatia, Greece, North Macedonia, and Serbia, which we studied in conjunction with those that preceded them in the Balkans and with published data from present-day people genotyped on the Human Origins array (34, 39) (Fig. 5). The reduction of Anatolian Neolithic ancestry was a long-term process in Southeastern Europe (1), which allows us to differentiate present-day populations from those preceding the Slavic migrations. When we order individuals along this component of ancestry (Fig. 5), we observe that present-day Slavs outside the Balkans have the least, whereas pre-Slavic inhabitants from the Balkans have the most of this type of ancestry, with present-day people from Southeastern Europe intermediate between the two extremes. Three individuals from Bulgaria (Samovodene), North Macedonia (Bitola), and an outlier individual from Trogir in Croatia (700 to 1100 CE) have the lowest levels of this ancestry. Most individuals from Trogir (a port city of the Adriatic in Croatia that was founded by Ancient Greek colonists and was part of the Byzantine Empire) overlapped with present-day people from ~700 to 900 CE, as did 12th century CE individuals from Veliko Tarnovo and Ryahovets in Bulgaria and a mid–fourth century CE Roman-era individual from Marathon in Greece, who, however, lacked the Balkan hunter-gatherer ancestry found consistently in the present-day population (Fig. 1). Finally, three medieval individuals from Albania (500 to 1100 CE) and a Late Antique (~500 CE) individual from Boyanovo in Bulgaria preceding the Slavic migrations, overlapped with the more ancient population, having high levels of Anatolian Neolithic ancestry. Among present-day people, Greeks and Albanians have more Anatolian Neolithic ancestry than their South Slavic neighbors. Slavic migrations have some echoes, ~3000 years later, to the spread of the descendants of Yamnaya steppe pastoralists into Southeastern Europe (1, 7). Although both events were transformative, any analogy should not be pushed too far. The medieval movements were carried out by large, organized communities engaging with complex states, such as the Avar Khaganate and Byzantine Empire, and no comparable polities existed in Yamnaya times. Collectively, our data suggest that although Balkan groups experienced a shift of ancestry in the medieval period, the fusion of locals and migrants was variable with individuals of diverse ancestry being present in medieval times and persisting up to the present."

Lazaridis et al. 2022

According to a thread "Southeast Europe/Balkans: From Neolithic to Modern times" made by Anthrogenica forum user Dorkymon in 2019, see image below, the first group was made by three Early Slavic samples (Bohemian and two Avaro-Slavic from Hungary) and the second group by three Bronze Age samples from current Croatian territory. They were compared with contemporary South Slavic samples, with a medieval Slavic estimation for Croatian samples ranging from 50 to 80%, usually around 70%, while other near nations of Serbia, Montenegro, Albania, North Macedonia, Bulgaria, Romania, and Moldova had an estimation of 50% or lower.

Image by user Dorkymon in 2019

Different calculator models using ancient & medieval or contemporary population samples at Vahaduo (Eurogenes K13), take into account that models with lower distances are more precise (~1.0 or <1.0), practically show the same thing.

Vahaduo Eurogenes K13 (Updated and Ancient), models with lower distances (on the right) are more precise, 05-2021

As observed in the recent Anthrogenica thread "New Samples from Migration Era and Early Medieval Moravia", Proto-Slavs had a homogeneous genetic profile distinctive from Germanic or Romance populations which can be seen even today because Early Slavic archaeogenetic samples cluster with modern-day North Ukrainians, Belarusians, and Lithuanians. They probably were an isolated and initially smaller population that rapidly exploded in numbers and expanded from their Urheimat. The proportion of the Slavic ancestry in contemporary Slavic ethnic groups in Central and Southeastern Europe shows that the regions settled by the Slavs in the Early Middle Ages mostly had a lack of native people and low population densities.

Made by Waldemar

Genetic diversity of the early Slavs (7.5.2021.) at Slavic origins blog

Made by Ph2ter
Made in June 2022 by Ph2ter

Graphic visualization of newest archaeogenetic samples' admixture, per post June 2022 by Ph2ter
 
In conclusion, according to the autosomal DNA scientific research, a medieval migration from Eastern and Central Europe to Southeastern Europe did happen, it was substantially large to influence the gene pool partly because those regions had low population densities, and depending on the Southeastern European country replaced approximately 30-80% of the native population ancestry, more specifically, the medieval Slavic component over time replaced approximately 30-65% of the native population ancestry. Throughout history, there existed different historical and archaeological theories that ranged from autochthonous to Pan-Slavic, culture-historical to culture-social but advancing or physical or cultural replacement both were wrong as overestimated or underestimated specific ancestral components or cultural influences in the ethnogenesis of those ethnic groups depending on the region. However, the medieval expansion from Eastern Europe which correlated and even caused population and cultural replacement is generally an undeniable fact which cannot be refuted or ignored anymore.

2. Y-DNA:

According to A global analysis of Y-chromosomal haplotype diversity for 23 STR loci by Purps et al. 2014, Hungary (143) belonged to I1 6.99%, to I2a1 15.38%, to R1a 23.78% and to R1b 24.48%, which together make over 70%. According to Origins, admixture and founder lineages in European Roma by Martinez-Cruz et al. 2016, Hungary (187) belonged to I1 6.41%, to I2a1 16.04%, to R1a 26.74% and to R1b 17.61%, which together make over 65%. According to the Croatian national reference Y-STR haplotype database by Mršić et al. 2012, in Croatia (1,100) belonged to I1 5.8%, to I2a1 37.7%, to R1a 22.1% and to R1b 7.9%, which together make over 70%.

According to High levels of Paleolithic Y-chromosome lineages characterize Serbia by Regueiro et al. 2012, in Serbia (103) belonged to I1 7.8%, to I2a1 29,1%, to R1a 20.4% and to R1b 7.8%, which together make over 60%. According to Šehović et al. (2018), which analyzed 179 samples from Mirabal et al. (2010), they belonged to I1 5.8%, to I2a1 39.8%, to R1a 14.6% and to R1b 4.8%, which together also make over 60%. According to Kačar et al. (2019), which analyzed 303 samples of Serbs in Serbia and near countries, belonged to I1 7.6%, I2 was represented by I-PH908 (25.08%), I2a1b3-L621 (7.59%), I2-CTS10228 (3.63%) and I2-M223 (0.33%), R1a was represented by R1a (10.89%), R1a-M458 (2.31%), R1a-YP4278 (1.32%), and R1a-Y2613 (0.33%), while to R1b 5%, which together also make over 60%. According to Y-Chromosome Diversity in Modern Bulgarians: New Clues about Their Ancestry by Karachanak et al. 2013, in Bulgaria (808) belonged to I1 4.3%, to I2a1 21.9%, to R1a 17.6% and to R1b 10.9%, which together make over 50%.

However, as it is stated at Eupedia (among others) on R1a and I2, the first haplogroup is almost exclusively represented by R-Z282 > R-Z280 > R-CTS1211/M558 and R-Z282 > R-PF6155 > R-M458 subclades among the Yugoslavs with the most probable expansion from Ukraine, Poland, and Czechia-Slovakia. While the Croatians almost exclusively fall into R-M558 subclades, almost equally both R-Z280 and R-M458 subclades are among Serbians, Bulgarians, and Romanians, which TMRCA and distribution "appears to be of Slavic origin". The R-M558 subclade is more frequent among East Slavs in Eastern Europe and Volga-Ural region, while R-M458 among West Slavs (especially Czechs and even East Germans) in Central and Eastern Europe (Underhill et al. 2015). The second haplogroup, also known as I2a-Dinaric, is mainly represented by I-CTS10228 > I-Y3120 including I-S17250 > I-PH908 (Dinaric South) subclades which had a very rapid formation and expansion since 1900-1700 YBP, "concordant with the timing of the Slavic ethnogenesis, considering that it takes a few centuries before one man can have enough male descendants to start having an impact at the scale of a population", and that "the higher percentage of I2a-Din in the south is probably just due to another founder effect due to the fact that the South Slavs originated in western Ukraine, where the ratio of I2a to R1a was higher".

Image from "Haplogroup R1a" (March 2018) by Vayda

Isofrequency contour map for haplogroup I2-CTS10228 (April 2021) by LP. See the full resolution.

In the end, if the autosomal percentages for Hungary, Croatia, Serbia, and Bulgaria as reference populations are compared to Y-DNA percentages of respective populations, with R1a and I2 subclades considered to be from the medieval migratory period and with some additional Celtic-Gothic or late medieval remains of R1b and I1 subbranches, forming at least 50-70% with south-north gradation, it results with a very positive or almost identical correlation to the autosomal 45-65% medieval migratory ancestry with south-north gradation.


2.1 I-PH908:

Also named as I2a1a2b1a1a1c at ISOGG phylogenetic tree from 31 March 2019, according to Bernie Cullen & Zdenko Marković, administrators of I-P37.2 haplogroup research project at genetic genealogy company Family Tree DNA, the SNP PH908 was first reported by geneticist Pille Hallast from Estonian Biocentre in 2014 (see a post from 22 August and post from 23 September 2016), as well such information is reported by GeneticHomeland. This one SNP represents >99% of the so-called Dinaric South or Balkan STR cluster, characterized by STR marker results DYS448=19 as well DYS449=30 which are reported since 2007 by Ken Nordtvedt, a cluster noted for "recent and rapid expansion over a wide geographical area" (see a post from 7 November 2016). According to YFull YTree (v9.02.00), the SNP was formed around 1950 YBP (Cl 95% 2200-1750 YBP) and had time to most recent common ancestor around 1700 YBP (Cl 95% 1850-1550 YBP) when emerged numerous subclades, many still not defined, indicating a rapid expansion before and during Early Middle Ages. For now, they are supposedly divided by STR DYS561=16/15 marker results making two large sub-branches, and the first has 9 while the second 2 immediately known subclades (see a post from 26 January 2021). According to Cullen (see a post from 3 April 2019), one scenario is that this, possibly elite individual, had many surviving sons and "each of these sons founded a male line that has survived until the present day" or more probably "had fewer sons with surviving male lines and that some of these lineages are from expansions in the second, third or fourth generation".

While the Dinaric South cluster is prevalent among Yugoslavs, the Dinaric North subclades are also significantly present among Slovenians, Macedonians, Bulgarians and Romanians. Roughly, 60-80% of I-CTS10228 > I-Y3120+ frequency in Western Balkans and 50% in Eastern Balkans is represented only by a single young subclade, I-PH908 (per scientific data, the analysis and results see blog post from November 2022). It shows that "despite haplogroup I-M423 reaches highest frequencies in Dinaric Alps region (Western Balkans), that is also region with it's smallest variation ... is one of many reasons why Western Balkans can not be place where I-M423 originated, nor it's descendant Slavic-Carpathian subclade" (post May 2020 by NevGen). Prior to the current knowledge of existence of SNP PH908, the haplotype clusters evolution and detalied development of the phylogenetic tree, as previously said, older studies like Šarac et al. 2016 erroneously claimed things like "high haplotype diversity of this lineage [I2a1b-M423] in Croatia reveals its significant expansion only after the adoption of agriculture by Mesolithic hunter-gatherers in SEE", not knowing that almost all examined haplotypes belong to I-CTS10228 > I-Y3120+ and vast majority (43/57) of them belong to a single subclade, I-PH908 (DYS448=19/18), hence revealing no relation to Mesolithic or Neolithic and SEE area and low haplotype diversity & lineages of both I2 and I-Y3120 and actually revealing high haplotype diversity of I-PH908 lineage.

I-PH908 Draft Tree (26-01-2021) by I2a Project

According to a doctoral thesis by geneticist Olga Utevska (2017),  Генофонд українців за різними системами генетичних маркерів: походження і місце на європейському генетичному просторі (transl. "The gene pool of Ukrainians revealed by different systems of genetic markers: the origin and statement in Europe"), the highest diversity of derivative Balkan cluster (DYS448=19) and ancestral Carpathian-Dniester cluster (DYS448=20) is in Western Ukraine, South-Eastern Poland, and Belarus, which is indicative of a place of origin and medieval migration route.

Frequency (left) and Variance (right), Utevska 2017

According to the blog post "DinA3, subclade PH908" (December 2017) by Vayda, it has approximately 9.3 million carriers of which 1/3 in Ukraine and Russia alone. Other visual attempts of contemporary and medieval placement of I-PH908 include:

Image from "DinA, subclade S17250" (December 2017) by Vayda
Image made by Anđelko Đermek (2017)

3 Origin place, culture, and migration of the Slavs:

Initially, the paternal haplogroup I2a1 was argued to have been a "major lineage of the Cucuteni-Trypillian culture and it was present in the Baden culture of the Chalcolithic Carpathian Basin" (Lipson et al. 2017, Neparáczki et al. 2019) some 4800-3000 BCE in Eastern Europe and 3600-2800 BCE in Central-Eastern Europe. However, that is not the case as scientific research until now mainly found G2a (Mathieson et al. 2018, Gelabert et al. 2022, Patterson et al. 2022), and like in other Southeastern European ancient samples only the I2a1a1-CTS595 or I2a2 subclades which are not of I2a1b2-L621 lineage which "formed 11500 YBP, TMRCA 6500 YBP" (YFull YTree). Although it is dominant among the modern Slavic peoples on the territory of the former Balkan provinces of the Roman Empire, I2-L621 lineage was not found in samples from the Roman period and is almost absent in the contemporary population of Italy (Fóthi et al. 2020). 

According to Mathieson et al. 2018 and Patterson et al. 2022, out of twelve male samples from Early & Middle Bronze Age and Early Iron Age Croatia, mostly related to Iapodes and Liburni tribe of Illyrians, almost all belonged to J2-L283 with the exception of two R1b and one I-Y3120. However, the I-Y3120 sample's (I18719 from Bezdanjača Cave dated 1500-1000 BCE) raw DNA file is not of good quality and reliable (proved by the staff of NevGen Predictor and others). It also has Y-DNA and mtDNA haplogroup (specifically subclade I-Y3120 which TMRCA doesn't correspond, and more importantly, HV0a1a1b with both recent 275 YBP and ancient 4400 YBP TMRCA matches in Germany and Northeastern Europe) as well as atDNA ADMIXTURE combination not found in any contemporary ancient sample from Croatia and Southeastern Europe. By atDNA it looks like a modern period sample as its autosomal genetic combination is of modern South Slavs which is not found in any other ancient sample. The specific archaeological sample has no carbon dating and is found in a cave with skeletal remains from the ancient up to modern periods. Most probably it is not a fault of contamination but misdated skeletal remains. This is additionally supported by many male Bronze & Iron Age samples from Slovenia, Serbia, and Hungary from the same studies, none of which carried the I-Y3120 lineage or had the same atDNA admixture of modern South Slavs. According to Olalde et al. (2021; published 2024), none out of 45 male samples from Croatia and Serbia dated 70-538 CE carried the I-Y3120 lineage, while three I-Y3120 lineage samples emerged only in the Serbian medieval period. According to Antonio et al. Stable population structure in Europe since the Iron Age, despite high mobility (2022; published 2024), also none out of 35 samples from Western Balkans dated 23-639 CE carried Slavic I2-L621 or R1a lineage nor showed (Balto-)Slavic atDNA admixture. According to Lazaridis et al. The genetic history of the Southern Arc: A bridge between West Asia and Europe (2022), besides two I-CTS10228 > I-Y3120 medieval samples from Croatia and Turkey, three Bronze Age I-L621 / I-CTS4002 / I-CTS10228 > I-Y3120 samples were found in Northeastern Romania, Bulgaria, and Croatia, however, all three yet remain to be thoroughly analyzed, are once again without carbon dating, Croatian sample (I18721) is once again from the Bezdanjača Cave and autosomally again [impossibly] closest to modern South Slavs but somehow archaeologically 1000 years older than I-Y3120 TMRCA, besides, other 37 Bronze Age & Iron Age samples from the same Cave and other Croatian sites almost exclusively again belonged to J2b and R1b subclades. According to Zagorc et al. Bioarchaeological Perspectives on Late Antiquity in Dalmatia: Paleogenetic, Dietary, and Population Studies of the Hvar Radošević burial site (2024), on an island where today Y-DNA haplogroups I2 and R1a together make 60-75%, none of the 10 male samples dated from 3rd to 5th century belonged to these haplogroups. Considering the number of over 150 Bronze & Iron Age ancient DNA male samples from many Balkan countries from these studies giving a frequency of I-L621+ samples around 0-1%, the pattern is obvious, it would be a very, very rare and atypical subclade for the Bronze & Iron Age time periods in the Balkans.

Even if presumably very rare "autochthonous" carriers could suddenly reproduce in sufficiently big numbers to explain the post-medieval & modern frequency, diversity, distribution as well as atDNA admixture, it would be logically and statistically impossible. Eventually, if the two Bezdanjača Cave samples dating is proven right after further ancient DNA sampling, carbon dating and Temporal population structure (TPS) method calibration - theoretically far-fetching, they could be traced only to some Northern-Eastern European small population that emigrated and their members temporarily lived near (Proto-)Celtic Hallstatt-La Tene cultures until perished in Southeastern Europe, and such Y-DNA, mtDNA and atDNA became present in Southeastern Europe yet again only since medieval migration period with Slavic migrations from Eastern Europe. In short, among the countless archaeogenetic samples of different periods and cultures in the area of Eastern and especially Southeastern Europe until the end of antiquity, there is not a single and reliable pre-medieval archaeogenetic sample that belongs to the young Slavic I2-L621>CTS10228>Y3120 or had atDNA admixture found in early and modern South Slavs.

By 2024 such prediction was confirmed. In the Ringauer et al. ancIBD - Screening for identity by descent segments in human ancient DNA (2023), the Bezdanjača Cave I-Y3120 samples (I18719 and I18721) per IBD do not match with Bronze Age samples from the same cave and others from the same period, which are all grouped together, but with medieval samples clustered in "Baltic" group proving these two samples are seriously misdated samples who are not from pre-medieval period. According to Olalde et al. 2023, which made an analysis of many ancient and medieval samples, concluded that in the big data set Y-DNA haplogroups I2a-L621 and R1a-Z282 are absent in the antiquity and appear 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".

According to Rębała et al. Y-STR variation among Slavs: evidence for the Slavic homeland in the middle Dnieper basin (2007), the comparison of interpopulation Y-STR haplotypes excluded "a significant contribution of ancient tribes inhabiting present-day Poland to the gene pool of Eastern and Southern Slavs, and suggest that the Slavic expansion started from present-day Ukraine, thus supporting the hypothesis that places the earliest known homeland of Slavs in the basin of the middle Dnieper". According to Fóthi et al. (2020), the distribution of ancestral subclades like I-CTS10228 among contemporary carriers indicates a rapid expansion from Southeastern Poland, which is mainly related to the Slavs, and the "largest demographic explosion occurred in the Balkans".

Until now I-Y3120 was found at the archaeological site Ilıpınar where were settled early medieval Asia Minor Slavs in the 7-9th century (I10430); Sunghir (sample Sunghir 6) from Early Medieval Russia (~900 YBP) which belonged to Y3120 > S17250 > Y5596 > Z16971 > Y5595 > A16681 (formed circa 1650 YBP); in many samples from the Hungarian conqueror period particularly the Hungarian elite (895-mid 10th century) which were positive to I-L621, including I-S17250 and a I-PH908 > FT16449 > MF2888 sample SP-10 ERS9945221 (Neparáczki et al. 2019, Fóthi et al. 2020, Maróti et al. 2022), "very often accompanied by Asian maternal lineages, indicating that I2a1a2b1a1a could be more typical for the immigrants than to the local population". The sample ERS9945221 is first SNP confirmed I-PH908 archaeogenetic sample; the I-Y3120 was found in 1 sample according to preliminary DNA results from Early Medieval Poland (900-1200 CE); as well I-CTS10228 and I-Y3120 subclades were confirmed in two Viking samples from Sweden (VK53) and Ukraine (VK542) with predominantly Slavic autosomal ancestry of which the second belonged to Gleb Svyatoslavich (11th century) who was a descendant of Rurik dynasty; two samples dating from the 14th to 15th centuries AD Bosnia and Herzegovina belonged to I-PH908 and so on. 

By 2024, new archaeogenetic samples in Central and Southeastern Europe continue to confirm such a conclusion. Interestingly enough, the distinctives of the I2-L621>CTS10228>Y3120 and some R1a lineages with the early Slavs is seen in their absence, especially of I-Y3120, among archaeogenetic samples of Germanic and Sarmatian tribes previously living in the territories of Central and Eastern Europe settled by the Slavs (Amorim et al. 2018Stolarek et al. 2023Carrión et al. 2024Schütz et al. 2024).

As can be seen at the YFull YTree (v9.02.00), the contemporary distribution of I-M423 clades and subclades I-FGC41353 > I-Y3104 > I-L621 > I-CTS10936 > I-S19848 > I-CTS4002 up till I-CTS10228 is in Western and Central Europe. It indicates that I2-L621 archaeological homeland in ancient times was not in Eastern neither Southeastern Europe - if excluded possibility of circular movement and emigration from those parts of Europe to the West in ancient time - and considering I-CTS10228's age estimation for formation at 5100 YBP (3080 BCE) and time to most recent common ancestor at 3400 YBP (1380 BCE), as well as the fact its subclades represent almost all percentage of I2 in Southeastern Europe, the old hypothesis about the continuous autochthonous origin of the haplogroup I2 among today's Balkan people and relationship to the Cucuteni-Trypillian & Baden cultures is completely debunked, wrong and out of question. The distribution of clades correlates with the Unetice > Tumulus > Urnfield culture (c. 1300 BC – c. 750 BC). Somewhere around the time of Balto-Slavic divergence and formation of the Pre-Slavic period (c. 1500 to 1000 BC), the I-CTS10228 > I-Y3120 lineage of Proto-Celtic origin from Central Europe moved to Eastern Europe and was assimilated into the Proto-Slavic population. In the following centuries, it was probably present in Milograd or Chernoles culture. In the process of Proto-Slavic ethnogenesis, it experienced steady demographic growth, becoming one of the dominant and characteristic Slavic Y-DNA haplogroups.

Map of the Urnfield Culture c. 1300 BCE by Simeon Netchev at WHE

Wikimedia Commons

There still needs to be found a contemporary or archaeogenetic sample which will make I-Y3120's time to the most recent common ancestor higher because the current is at 2100 YBP (1st century BCE). It has at least five subclades, I-FT76511, I-Y18331, I-Z17855, I-Y4460, and I-S17250 out of which the I-S17250 is the most numerous. The archaeological cultures in Eastern Europe had a heterogeneous and complex interaction. Considering previous information and maps which placed the subclade around Southeastern Poland and Western Ukraine, there exist two options:

1) Przeworsk-Zarubintsy cultures (3rd century BCE - 2nd century CE), particularly Zarubintsy in the East, which are considered to be related to Early Slavs, but with some Celtic, Germanic, Scytho-Sarmatian, and Dacian influences. The latter was replaced by Kyiv and Chernyakhov culture (2nd century CE - 5th century CE), which will be replaced by late 5th and mid-6th-century Slavic cultures of KolochinPenkovkaKorchak, and Ipotesti–Candesti.

Wikimedia Commons
2) Lipitsa culture of Dacian tribes primarily of Costoboci, who forcibly moved near tribes of Carpi and then formed Carpathian Tumuli culture between the end of the 2nd and end of the 4th century CE. It will be replaced by Chernyakhov culture, but Korchak culture kept the tumuli-kurgan type of burial.

Wikimedia Commons
The regional ancient tribes of Bastarnae, Costoboci, Carpi, Daci, and others were numerous, but although they did have expansions and had a lasting war with the Roman Empire in the 3rd century, at the same time it was their downfall. They lost and were relocated by the Romans around the Danube or else as laeti or to defend the limes, with a substantial part of them being Romanized or assimilated by 4th century Gothic and Hunnic hordes who temporarily ruled the region. Considering also their limited Southern localization it becomes a bit difficult to argue whether from such a population emerged a rapid, numerous, and widespread expansion which will be vital to the following Proto-Slavic culture to the Northwest and Northeast. Perhaps it could be possible if the descendants of the common ancestor regularly moved, changed their ethnic and social identity, and managed to successfully use new political movements for their own reproduction, being swapped into Hunnic-Gothic and then Slavic expansion, but that for now remains difficult to argue.


Büntgen et al. "Cooling and societal change during the Late Antique Little Ice Age from 536 to around 660 AD", (Nature, 2016)

Interestingly the I-Y3120 sub-subclades date of formation and growth in Eastern Europe happened around the time of Antonine (165-180 CE) and Cyprian (249-262 CE) plagues and so-called Roman Warm Period with increased climate variability with periods of cooling and drought (3rd-4th century CE), while subsequent great spatial expansion with the Slavs happened during the Justinian (541-542 CE) and Roman (590 CE) plagues and extreme weather events of 535–536 and Late Antique Little Ice Age (536-660 CE). The epidemics, climate change, weather and crops, population pressure, "primeval urge" to push into the Mediterranean, "domino effect" of tribes forcibly moving westward in Eurasia, Roman Empire's political, cultural, and economic falling apart positively or negatively influenced both the "barbaric" tribes and Roman Empire's population and shaped their mutual relations during the migration period (c. 300-800 CE).

The approximate date of I-PH908's time to the most recent common ancestor (1700 YBP), 320 CE, coincides with the upcoming Slavic movement. Most probably it will be never known the tribe of origin and by that time most probably was not part of only one Slavic archaeological culture. The Slavic movement began North of the Danube, and according to J. P. Mallory's In Search of the Indo-Europeans: Language, Archaeology and Myth (1989), their origin was between the Dniester and Vistula rivers where from at least 1500 BCE until the historical appearance of the Slavs can be traced a cultural continuity. The most plausible candidate would be Vistula Veneti, an umbrella term for a population considered to have been ancestral to newly emerging Wends (West Slavs), Sclaveni (Central-East Slavs), and Antes (East Slavs). Considering the contemporary wide distribution of I-Y3120 (> I-PH908) subclades the most plausible archaeological culture in which emerged had to be somewhere in the East, but not too far, not in the West or South, somewhere that Slavic migrations happened from East to the North and West as well later to the South. According to Irena Cvijanović's The Typology of Early Medieval Settlements in Bogemia, Poland and Russia (2013), it is assumed that the "settlements of the Prague-type culture are widespread in Bohemia, Southwestern Slovakia, and Poland ... originated in the steppes of Western Ukraine and that it appeared in the fifth century", and Zdeněk Váňa's The world of the ancient Slavs (1983), "the first archaeological traces of the historical Slavs were finds of simple, mostly undecorated pottery ... between the eastern Carpathians and the rivers Pripyať and middle Dnieper" known as Korchak type, as well "the archaeological finds of the Prague type document the advance of Slav tribes from the east to the west", hence the most plausible would be Korchak culture of Western Ukraine.

Váňa 1983

Proto-Slavic archaeological cultures in the beginning of the Middle Ages with Prague-Korchak (/), Penkovka (\), their transition and mixing (X) which influenced the creation of the Ipotesti-Candesti (dots) culture 

According to Peter Heather's Empires and Barbarians: The Fall of Rome and the Birth of Europe (2009), "on the basis of Jordanes’ report of the relative geographical distributions of the Sclavenes and Antae, Penkovka has often been thought to have been the product of the latter, and Korchak of the former. These precise identifications are questionable, but the basic similarities of the two systems, combined with the geographical coincidence between their spread and where we actually find Slavic speakers in the sixth century, does make it reasonable to think of Penkovka, like Korchak, as at least Slavic dominated".

3.1 Slavic tribes related to Korchak culture:

To confirm the hypothesis of the connection with Korchak culture will be examined tribes related to the Korchak culture. There exist many medieval Slavic tribes who were related to the Korchak and other cultures and it is difficult to choose any one of them as mainly I-PH908 carriers. Most probably most of the Slavic tribes and migration waves had some higher or lower percentage of haplogroup I-Y3120 and hence I-PH908. Considering the cultural location, migration, and distribution of I-PH908 candidate tribes probably had to be those who were big enough and at the same time present among both East and West Slavs to influence their ethnogenesis. In addition, tribes who migrated to Southeastern Europe - such a criterion corresponds to two candidates - the White Croats and Dulebes. According to V. V. Sedov's Slavi︠a︡ne v rannem Srednevekovʹe (1995) the latter is considered to have been a Proto-Slavic tribal union whose tribes lived in Western Ukraine, the Southern Czech Republic, and the Middle Danube area between Lake Balaton and the Mur River in Hungary, but some hydronyms and toponyms can be found also in the Western Balkans, besides that, there is anything much to say.

Croats (X within \) and Dulebes tribes (Volhynians, Drevlians, Polans within /), Sedov 1995
Croats and Dulebes and their presumable migration routes are within the range of Prague-Penkovka culture, Sedov 1979
According to Sedov (1995), the White Croatian tribes emerged from Antae Penkovka culture and from there emigrated to the West and divided into several groups who settled in different Early Slavic territories. A. V. Majorov in Velikaja Horvatija: etnogenez i rannjaja istorija slavjan Prikarpatskogo regiona (2006) argued the territory which White Croats inhabited in the Middle and Upper Dniester and the Upper Vistula river was also part of Korchak culture related to Sclaveni which was characteristic for the Kurgan-type of burial which was also found in the upper Elbe territory where lived the Czech Croats. According to Cvijanović (2013) in Western Ukraine and Southeastern Poland was located Korchak-Penkovka culture borderland from which Early Slavs emigrated North, West, and South. Later in the 8th and 9th centuries, they developed the Luka-Raikovetska culture to which belonged Croats, Buzhans, DrevliansEastern Polans, TivertsiUlichs, and Volhynians. According to Vera Gupalo's Звенигород і Звенигородська земля у ХІ–ХІІІ століттях (2014) and Mykhailo Kuchynko's Літописні хорвати: проблема етноплемінної належності та політичної залежності (2014), archaeological data shows a common tribal origin of the Prague (6-7th) and Luka-Raikovetska (8-10th century) culture in the territory of the Eastern Carpathians which is roughly defining the territory of the Carpathian Croats by the Carpathian ridges on the West, the Khotyn and Chernivtsi in the Southeast, the watersheds of the Western Bug and the Dniester river in the North, and within it, they had extraordinary large gords like Plisnesk (450 ha) and Stilsko (240 ha) among others and each could accommodate tens of thousands of people indicating a very large and well-organized population.
Both Łowmiański (1964), Sedov (1995), and Majorov (2006) with many other scholars agree that they were a large Proto-Slavic tribe or among the oldest Slavic tribes which got scattered in various directions during the Slavic migration period. According to Łowmiański, there's little doubt that a large group of Croats lived in Lesser Poland and that the tribes of Vistulans and Lendians (Lyakhs) were actually tribes of Croats or Croatian tribal alliance. According to Váňa (1983), in the distant past their territory "stretched to the north of the Carpathians, today's southern Poland and western Ukraine, from where the mighty Slav tribe of the Croats spread, some to the south, the Balkans, others to the west as far as eastern Bohemia". According to historical sources, White Croats in the 10th century were in friendly and matrimonial relations with the Hungarians and Árpád dynasty.

According to Croatian origo gentis five brothers and two sisters (7) led the people to the South; which slightly reminds of Bulgar Kubrat's five sons of whom Asparuh subjugated Seven Slavic tribes (7); seven tribes and chieftains of Hungarian-Magyars (7); 10th-century L'Abrege des Merveilles account about a nation between the Slavs and the Franks which follows the religion of the Sabeans and worships the planets, is a very intelligent and skilled in the arts of war, making war on the Slavs, the Burgan, and the Turks, and it has 7 feasts in the year, according to the name of the 7 planets, and that of the Sun is the most splendid; 13th-century chronicle about seven (7) or eight Gothic-Slavic tribes who arrived in Croatia during mid-6th century. They are also rich in legendary stories, from 10th-century epic poem Widsith and 13th-century saga Hervarar saga ok Heiðreks which possibly mention them near river Vistula or the Carpathian Mountains, to 12th-century Ukrainian-Russian legendary story about brothers Kyi, Shchek and Khoryv of whom the latter is by some scholars related to Croatian ethnonym, 14th century Czech and Polish legendary story about brothers Lech, Czech, and Rus of whom the first two came from homeland "Croatia" as Váňa (1983) concludes "a reference to an older folk lore, it cannot mean modern Croatia, as Dalimil himself must have assumed, but Great or White Croatia" and that legendary Czech came with six brothers (7), but notes that "it is thus possible to cull some truth from legends. But they cannot be taken as providing present-day historians with reliable facts".

Black dots are Kurgan burials in Czechia, Croats (X), Dulebes (|), Moravians (\\), Sedov 1995

In a map (see below) with red color are designated boundaries of the territory on which presumably lived White Croatian tribes according to multiple historians and archaeologists, with noted several gords and toponyms with Croatian ethnonym. In yellow color is lined the boundary of the territory of Lusatia where live the contemporary Sorbs, sometimes called the Lusatian Serbs (more about them below), while in green color is lined the presumed boundary of the Serbian tribe led by Dervan in the 7th century. With blue color lines and arrows are designated points of skeletal/cranial excavation sites which were researched by craniometric methods. As a reference was used work by M. Šlaus Craniometric relationships of medieval Central European populations: Implications for Croat ethnogenesis (1998) and Kraniometrijska analiza srednjovjekovnih nalazišta središnje Europe: Novi dokazi o ekspanziji hrvatskih populacija tijekom 10.do 13. stoljeća (2000). According to it, Old Croatian sites from Dalmatia (Nin, Danilo, Mravnice, Bribir) and two (Bugojno, Gomjanica) from Bosnia and Herzegovina, belong to the cluster with Old Polish sites (Cedynia, Wišlica, Lednicki), and on study's diagram, Nin and Cedynia in parameters are basically identical. However, on the map is taken Lednicki as the starting point of the migration because it is most Southern Polish site, and because in the second research paper is considered the direct route through sites Nitra-Lupka -> Zalaszabar-Dezsosziget -> Nin-Ždrijac. With black color lines and arrows are designated points of skeletal/cranial excavation sites which were researched by craniometric methods, and as reference was used anthropological work by T. A. Rudych of medieval sites of Halych (2011) and Zelenche (2015) as well as older Ukrainian-Russian anthropological literature which found closeness between the medieval Galician and medieval Croatian populations (as well as Czech, Moravian, and Lusatian) which "can testify for their common origin".

In the territory of Western Balkans, the Croatian population from the narrow littoral core expanded to the North, initially into Bosnia and Herzegovina by the 10th century, and then into Zagorje (Northern Croatia) and Slavonija (Eastern Croatia) after the 11th century, as according to research the initial skeletal remains of the Northern and Eastern Croatia do not belong to the Polish-Old Croatian cluster, but so-called Avaro-Slav cluster (roughly cluster of Slavic sites West/East of Danube). According to Bašić et al. Cultural inter-population differences do not reflect biological distances: an example of interdisciplinary analysis of populations from Eastern Adriatic coast (2015), medieval skeletal remains of Šopot and Ostrovica (also from Northern Dalmatia) also belonged to the Polish-Old Croatian cluster, concluding that "As previously shown (6), all the sites were in vicinity to Polish sites and were separated from the other analyzed sites. PCA showed that all Eastern Adriatic coast sites were closely related in cranial morphology, and thus, most likely had similar biological makeup".

For a high-quality map see Scribble Maps

The archaeogenetic Slavic sample I15742 of a "White Croat" woman from the Dalmatian city of Trogir who is radiocarbon dated to 661-775/677-709 (Lazaridis et al. 2022), according to the Ringauer et al. ancIBD - Screening for identity by descent segments in human ancient DNA (2023) is confirmed as a direct cousin of AV1 and AV2 Avaro-Slavic family samples from Szólád on southeastern coast of lake Balaton in Western Hungary dated to 549/560-640/645 (Amorin et al. 2018). The samples AV1 and AV2 are also related to I-Y3120 medieval Slavic samples of I10430 and Sunghir6 as well as many Avarian and Hungarian samples from Central-Eastern Hungary (and some Vikings showing their partial Slavic origin), while I15742 is predominantly related to the Hungarian elite and commoners from Eastern Hungary. Croatian sample I15742 also belongs to haplogroup H3h6 matching 9-10th century northeastern Hungarian commoner and Viking woman from Italy.

Early Croat I15742 sample from 7-8th century in Trogir is direct cousin of Avaro-Slavic samples AV1 and AV2. The Croatian sample's other Avar-Hungarian period matches cluster are in full line, while AV cluster is in broken line. It indicates that the migration(s) went from Northeastern direction.

It is considered that the White Croats were assimilated into Czech, Polish and Ukrainian ethnos, and are one of the predecessors of the Rusyns, as well as one of the ancestors of the contemporary Croatian nation. Although there is no ancestral ethnic group that preserved Croatian ethnic identity as a hypothetical representative group for comparison of Y-DNA haplogroup distribution will be considered Rusyns, the population of Western Ukraine, and the city of Kraków in Lesser Poland, who are considered in Ukrainian and Russian encyclopedias among other academic sources as primarily the descendants of White Croats (see Rusyns#Origins). According to Veselinovic et al. (2014), Pannonian Rusyns (200) from the Vojvodina region in Serbia are R1a 43%, of which at least half are R-CTS1211/M558 subclade, and around 20% have I2, of which less than 5% I-PH908. The mtDNA study by Nikitin et al. (2009) of their sub-groups Boykos, Hutsuls, and Lemkos revealed high intra-population differences, and Hutsuls clustered closest to the Croatian (0.11) and Ukrainian (0.16) population. The Western Ukraine studies sometimes do not have well-defined Y-DNA subclades but practically the whole I2 percentage in Ukraine is represented only by I-CTS10228 > I-Y3120, and according to Mielnik-Sikorska et al. (2013), the population of Lviv Oblast is 44.82% R1a of which R-M458 is 13.65%, and 28.57% I of which 2.5% is I-PH908, while according to Utevska et al. (2015) almost the same percentages of R1a and I2-P37.2 (~25-30%) are found in Zakarpattia Oblast, while in Chernivtsi Oblast the I2 with 35% occurs even more frequently than R1a-M198*. According to YHRD-YP000203, in the city of Kraków, R1a is almost 50% of which R-M458 is 32%, and around 11% is I-CTS10228 of which 3% is I-PH908. To be noted are also Hungarian Conqueror's period R1a and I2 samples in the Pannonian Basin, which are including the Hungarian elite 10th century sample ERS9945221 in Northeastern Hungary whose I-PH908>MF2888, located on the same route of migration on which passed White Croats through Pannonia to Dalmatia as well as those White Croats who remained in their ancestral homeland nearby had close relations at the time with the Hungarians, being only Slavs for which is said to have had such relations with the Hungarians and best candidates for person's origin.

In comparison, contemporary Croats under R1a (c. 20-25%; for more details see blog post from January 2023) mainly belong to subclades of R-M558 which is predominant (19.2%, firstly R-CTS3402>R-Y33>R-CTS8816>R-Y1392/Y2902>R-Y3226>R-Y3219>R-YP1144>R-PH3519>R-PH3782; secondly R-CTS3402>R-Y2613>R-Y2609>Y2608; thirdly R-YP343>R-YP340>R-P278.2>R-YP371; and fourthly R-CTS3402>Y33>CTS8816>Y3301>L1280; with a small minority of others), and R-M458 (4.9%, evenly divided betwen R-L1029 and R-L260>R-YP1337>R-Y132940), while R-Z282 is rare (1.2%), as well to I2 > I-CTS10228 > I-Y3120 (c. 30-35%), which is mostly represented by Dinaric-South cluster (I-PH908), while Dinaric-North cluster is higher only in North Croatia. The lack of determined subclades limits the possibility of assumptions and conclusions, but they mostly have matches in south-eastern Poland, western Ukraine, and Slovakia. It is expected that the Croatian tribes were more similar to the "ancestral" population and seemingly exist some correlations between contemporary populations.

R-Y2608 haplotype distribution (YHRD) 
R-PH3782 haplotype distribution (YHRD)
According to DAI and other primary historical sources, neighboring tribe to the Northwest of White Croats were the Serbs who also emigrated to the South, however, for them has been established in the scholarship to have lived around the region of Bohemia and Saxony, and unlike Croats, there is no proof that Serbs ever lived within Bohemia or in other parts of Europe (see White Serbia#Dispute).

Older archaeologists like Joachim Hermann since the 1960s argued that the medieval Sorbs would be a mixture of 6th-century Slavs with Tornow-type and 7th-century second wave Slavs of Leipzig-Rüssen type culture, with the latter influenced by middle Danube Avaro-Slav culture and to have brought the S(o)erbian language and identity as well as that the contemporary Upper Sorbian dialect which is spoken by the majority of contemporary Lusatian Sorbs is related to that culture. It practically claims that the Lusatian Sorbs by the 9th century adopted Serbian identity, but not only other scholars like Heinz Schuster-Šewc consider it completely unfounded, the contemporary archaeology since the 1980s also rejects such links because, as summarized by Mats Roslund's Guests in the House: Cultural Transmission between Slavs and Scandinavians 900 to 1300 AD (2007), they "have no solid basis in the written and archaeological evidence", previous datings of the ceramics were wrong by several centuries showing instead 8-13th century social stratification and Carolingian influence, and the old theorization was influenced by GDR's politics. In conclusion, as historian Heather (2009) explains, the "new chronologies have also put paid to older theories that an initial Slavic penetration into the Elbe region in the later fifth or sixth centuries was followed by a second wave of migration in the seventh. This hypothesis had in mind a potential parallel with the Serbs and Croats and the Balkans ... From all these materials, therefore, a clear enough picture emerges of a ribbon of Slavic settlement extending westwards from the northern hinterland of the Carpathian Mountains as far as the northern reaches of Slovakia in or around 500 AD. About fifty years later, a Korchak-type material culture penetrated south into the river valleys around the Middle Danube, and pushed on westwards to Bohemia. Another fifty years further on, and Slavic groups were both threatening the fringes of Bavaria and establishing themselves in the Elbe–Saale region".

It is often considered that their earliest mention is by Vibius Sequester who recorded them living on the other part of the river Elbe which divided them from the Suevi (Albis Germaniae Suevos a Cerveciis dividiit), with the district later recorded as Ciervisti, Zerbisti, and Kirvisti. The early mention some sources like Tadeusz Sulimirski's The Sarmatians (1970) interpret as remnants of the Alanic tribe of Serboi who after the fall of Huns remained behind retreating Germans and "ultimately merged with the subjugated Slavonic population". Nevertheless the possible connection with Serboi and Cerveciis, according to Joachim Herrmann's Slavs, Avars and the Merovingian kingdom (1996), the 7th-century Frankish source by Fredegar mentions Serbs in 630-631 under the leadership of Dervan and that they were for a long time ruled by the Frankish kingdom, indicating that they settled the Saale-Elbe valley at least since the late 6th century, almost the same is stated in DAI, and that unsuccessfully revolted against the Franks during the Samo's union. According to DAI's origo gentis when two brothers succeeded their father (probably Dervan or his close relative), one brother took "half of the folk" and "came as the refugee to Heraclius", being known as the "unknown archon" who founded the Serbian royal Vlastimirović dynasty. The migration to the Western Balkans most probably was related and influenced by the defeat against the Franks and it is widely regarded "that they possibly arrived as an organized military elite which managed to organize other already settled and more numerous Slavs".
Map of the Sorbian-Lusatian tribes between the 7th and 11th century by Wilhelm Boguslawski, 1861

Sorbs in the Northwestern dotted area while tight vertical lines correspond to the Middle Danube sites and later initial period of Greater Moravia, Sedov 1995
Creative Commons

Since the ancestral ethnic group survived and preserved its ethnic identity until today in Lusatia, hence contemporary Sorbians and Serbians share the ethnic identity, their population genetics comparison is very valuable to understand the Slavic migration. Considering the location of the medieval Serbian tribe in addition will be considered the East German population of Leipzig and other cities in Saxony.

In Lusatia: according to Veeramah et al. Genetic variation in the Sorbs of eastern Germany in the context of broader European genetic diversity (2011), "despite their geographical proximity to German speakers, the Sorbs showed greatest genetic similarity to Polish and Czech individuals ... showed evidence of subtle levels of genetic isolation", and according to Rębała et al. Contemporary paternal genetic landscape of Polish and German populations: from early medieval Slavic expansion to post-World War II resettlements (2013), Sorbs (123) who speak Upper Sorbian dialect predominantly 65% have R1a of which mainly is subclade R-M458 (56.9% and R-L260 is c. 41% of which majority is R-Y2905, while R-L1029 is c. 16%), and 4% have I2 but only 2.43% belong to I-CTS10228 of which 0% is I-PH908. In addition, almost 4.9% belong to the E-L540 cluster, 9.8% to I1, and 2.43% to J2 respectively.

In Saxony: according to Parker et al. A systematic investigation of human DNA preservation in medieval skeletons (2020), a study of eleven unrelated samples dated between the 11-14th century from an "abandoned medieval settlement of Krakauer Berg, near Peißen" in Saalekreis district in the state of Saxony-Anhalt and in the Early Middle Ages part of the territory of "Sorabos, called Colodici" (839 CE) of Kesigesburch/Kozinesburg located today at Zörbig in Anhalt-Bitterfeld district in the same state and distant less than 30 km: three male samples belonged to R-M458 > R-L1029 and one to I2-Y16473I1-Y4870E-L540 > A6295 and J2-Z1043 > BY38004 both of which seemingly have some rare matches from Serbia, while KRA009 sample whose R-L1029 > R-BY195372 on mtDNA subclade H13a1a1e has matches from Serbia and Albania. Most of the other KRA samples by mtDNA subclades seemingly also have some matches in Southeastern Europe. All of the samples "were observed to fall well within the expected [autosomal] variation of the Sorbs".

According to YHRD-YP000209 on 23 Y-STR markers in the city of Leipzig (303), 25% of people have R1a represented mainly with 13.53% by major subclade R-M458 (R-L260 is 4.62% and R-L1029 is 8.58%) and again only 3.97% belong to I-CTS10228 and 0.33% to I-PH908, making only 8.33% of I-CTS10228 percentage. In addition, have almost the same frequencies of E-V13, I1, and J2. According to manual analysis of another study for the same city (215) from Ballantyne et al. 2014 on 17 Y-STR, 20% of people have R1a and again only 3.72% belong to I-CTS10228 and 1.39% to I-PH908, making 37.5% of I-CTS10228 percentage. In addition, again have almost the same frequencies of E-V13, I1, and J2 as in Lusatian Sorbs. In total on more than representative 518 haplotypes for the city and region of Leipzig, R1a is 23.16%, I-CTS10228 is 3.86% while I-PH908 is 0.77% and making only 20% of I-CTS10228 percentage. 

The analysis of the frequency of I-L621 > I-CTS10228 on a very high number of 2179 samples with Minimal (9-loci) haplotype at YHRD from Leipzig (1114; YP000209), Chemnitz (743; YA003256), Halle (234; YA003081), and Dresden (88; YA003189) was manually checked by comparing 119 different I-CTS10228 haplotypes found in I2a FTDNA Project including all haplotypes from Germany, Czech Republic, and Poland. Few of them possibly can be found in non-I-CTS10228 clades, which is arguably positive because then the percentage will be higher or negative if the percentage is still very low. They consistently and firmly confirm that I-CTS10228 is barely 2% hence I-PH908 subclade is even less at around 0% in the contemporary population of Saxony. In conclusion, the genetic results clearly show genetic and anthropological continuity between the population of Saxony and Lusatia.

Calculation of I-L621 > I-CTS10228 frequency in Saxony using YHRD

In Northwestern Czechia: a fringe theory argues the existence of the so-called Naab Sorbs in eastern Bavaria and northwestern Czechia, although they are never recorded in historical sources. Some old I2a1 / I-Y3120 maps (1, 2) also wrongly show regionally elevated frequencies of 10-15% for I2-Y3120 in northwestern Czechia and beyond, making space for a lot of misconclusions. Northwestern Czechia is formed out of three regions, which sample size and frequency according to Zastera et al. (2010) is, from south to north, Plzeň (62; 8.06%), Karlovy Vary (31; 12.9%) and Ústí nad Labem (87; 8.04%). The study covered each of the 14 Czech regions, but Karlovy Vary is an exception both in terms of the percentage and the small number of samples. Only in the far east Moravian-Silesian exceeds 10% and all other regions have from 40 to 595 samples and a percentage below 8-9%, which means that the sample for Karlovy Vary was insufficient, which is confirmed by the percentages of the surrounding regions. If taken into account with or without the share of the population, the percentage obtained is between 8.9% and 9.6% in northwestern Czechia. According to Kushniarevich et al. 2015, Klatovy (48; 14.6%) in the Plzeň region has elevated frequency as well, but its population size is small, and when calculated with population distribution using only Plzeň's city population number, in the Plzeň's region the frequency is at a maximum of 9.4% and when that number is used to calculate the percentage of northwestern Czechia while using population size distribution gives 9.3% with all three regions and at maximum 10.63% with only two regions (Plzeň, Karlovy Vary).

If the scientifically outdated and rejected old archaeological thesis about the migration of the Serbian and other tribes from the Middle Danube valley to the Elbe river in the early 7th century is correct then the haplotypes found in Germany and the Czech Republic should derive from the supposedly ancestral samples in Serbia and not the other way around even if happened a subsequent migration to the Western Balkans. Also, considering that the migration of the Slavs to Saxony and Thuringia was around 600 CE which is somewhere between 1390-1450 years before the present, the age estimation of the formation and TMRCA of a subclade on YFull YTree has to be at the maximum of 1350 or fewer years before the present because a minimum of 10-15% additional age should be added to the YFull age estimation, especially if the samples are of an old NGS test as well as some reliable SNPs aren't taken into account of age estimation due to YFull criteria and so on, while a single SNP can greatly influence the age and disrupt the assumed connection between populations in question. There mustn't be any significant technical issues with age estimation. Unfortunately, that's exactly the case with matches at I-PH908 subclades like I-Z16983 > I-A8741,  I-FT16449 > I-MF2888 or I-Y32084 > I-A20333 where age estimations are based on samples tested at different companies by tests with different qualities and analysis done on different DNA files besides already excessive YTree age estimations which can only become bigger. They show a split in the Proto-Slavic homeland. It is unscientific and frivolous to take these results into serious consideration because besides sample bias since such samples are representing 0% in the contemporary population they currently most probably do not indicate a historical connection with the Serbian tribe at all but simply traces of the general movement of the Slavs and their archaeological cultures to the area of Saxony, Thuringia and all of Eastern Germany.

In comparison, contemporary Serbs with R1a (c. 15-20%) mainly belong to subclades of R-CTS1211/M558 (> R-L1280 and R-Y35>R-YP4278 among others) and a minority of R-M458 (> R-L1029 with a minority to R-L260 > R-Y2905 and R-A11460) and R-Z280 > R-Z92 (subclade R-Y4459>R-YP617>R-Y38448>R-BY169316), and they as well have a high frequency of I-Y3120 (c. 30-40%) which is mostly represented by I-PH908. The subclades R-M558>R-Y35>R-CTS3402>R-Y33>R-CTS8816>R-Y3301>R-L1280>R-Y5647 and R-M458>...>R-L1029>R-YP417>R-YP418>R-YP728>R-YP6047 and R-L260 > R-Y2905 are often present among Serbs in Serbia, Montenegro, and Bosnia and Herzegovina, and although based on YFull YTree can be argued that arrived from Eastern Europe, can be extensively found in both Central and Eastern Europe and as such cannot be completely excluded a possible migration of a part of both R-L260 and R-L1029 or some R-M558 subclades from Central Europe. However, the majority of R-M458 and R-M558 subclades they have also share with neighboring Bulgarians, Macedonians and Romanians where they are more prevalent indicating that such subclades are not good candidates for putative subclades of Serbian tribes.

R-L1280 haplotype distribution (YHRD)

R-YP418 haplotype distribution (YHRD)
Considering the historical facts it is expected that the Serbian tribes would have been more similar to the ancestral population and vice versa, but that is not the case with the contemporary Serbs and Sorbs and the general population of Saxony. Even if we suppose that the Serbian tribes in the minority brought I-CTS10228 > I-PH908 with them to the Balkans then the I-PH908 subclade or I-CTS10228 clade and especially macro-haplogroup I2 would have been present in more significant frequencies in contemporary Sorbs in Lusatia and the general population of Saxony. Not only they are not but are to the point of being almost non-existent. These results show that although they share an ethnic identity, happened an ethnogenetic discontinuity between these two ethnic groups and that almost all of I2 and R1a, possibly excluding rare cases of subclades of R-M458 and R-M558 and of other haplogroups (like R1b, E-V13, I1 and J2), among the contemporary Serbs originate from a different Slavic migratory route and population.

The reasons are many, from a small population of the migrating Serbian tribe which most probably arrived as an elite that non-biologically assimilated other already present and more numerous Slavs who accepted and continued to carry Serbian tribal identity due to political reasons similar to the case of Hungarians and Bulgarians. Previous settlement of those other numerous Slavic tribes in the territory of contemporary Serbia (Braničevci, Timočani, Moravljani, etc. possibly also part of the Seven Slavic tribes) which initially were not part of early medieval Serbia but First Bulgarian Empire and only since 12th century conquest started to be Serbianized. A long-distance between Serbian and Sorbian population and bottleneck and founder effects in both populations except that such a consistently very low frequency of I2 among Sorbs and other East German settlements cannot be sufficiently and reasonably explained with the distance and such anthropological processes. The firm consistency indicates the contrary - that the population in Saxony remained almost the same since the early medieval times and the Serbian tribe had the same genetics as them which was characterized by the lack of frequency of haplogroup I2. It is most probably related to the extensively detailed account from 10th DAI's 32nd chapter "Story of the Serbs" which reports about almost three generations of civil war in the Principality of Serbia and international war with the First Bulgarian Empire which lasted from mid-9th to the first half of the 10th century resulting in total depopulation because of which the Serbian principality was deserted for several years and then re-populated from neighboring countries Croatia, Bulgaria and others which was helped by Byzantine Empire. It is also indicative that according to the oral tradition recorded in DAI the family of Michael of Zahumlje was not of Serbian origin yet originated from the unbaptized inhabitants of the river Vistula called Litziki (Lendians) which geographically would relate them to the White Croats. As the Serbian tribes did not live and migrate to the Balkans from the Vistula valley and the Carpathians, and the ethnic accounts in DAI regarding the Sklavinies (South Slavic polities) population are a reflection of the political and not ethnological situation in the 10th and earlier century - including the 11th century contradictory accounts about the population of Dioclea by Byzantine historians not knowing difference between Croats and Serbs - it only further problematizes the claim about the uniform ancestry of the population in the supposed early medieval Serbian lands and throughout the centuries.

Quote of De Administrando Imperio and a map of the late 9th century with approximate borders of early medieval Principality of Serbia as well as projected borders (yellow) of modern day Serbia

In conclusion and contrary to previous assumptions, Korchak culture probably wasn't genetically uniform as the Slavs of Sukow-Dziedzice and westward Prague-Korchak-type culture, represented partly by Wends and Sclaveni, aren't good candidates for spreading I-Y3120, especially I-PH908, in Southeastern Europe. The Slavs who are most probably responsible belonged to the core and eastward Korchak-type and Penkovka-type cultures represented by the Sclaveni and Antae who moved from and along the Eastern Carpathians.

3.2 Slavic migrations to Southeastern Europe:

Although in the late 5th and early 6th century incursions of Germanic and Hunnic tribes possibly did happen first intrusions including some small groups of Sclaveni and Antae, the first sure incursions over the Roman limes started around 518 CE in the province of  Thrace followed soon in Illyricum almost every year for over a century and permanent settlement roughly began since 550 or 551. The devastating raids mainly began with the 560s arrival and leadership of the Pannonian Avars, culminating in sieges and destruction of Roman cities between 582 and 626 (Whitby 2017). The relations between Sclaveni, Antae, Bulgars, Avars, and Byzantines were complex and changed from hostile to ally depending on the political situation. Initially, Avars were used by the Byzantines to attack the Slavs, Antae temporarily were Roman allies, Sclaveni and Antae sometimes were at war, and so on.

F. Dvornik 1956

According to P. M. Barford's The Early Slavs: Culture and Society in Early Medieval Eastern Europe (2001), "the first half of the seventh century also saw the settlement of abandoned areas of the countryside of the Balkans by Slav families and communities. We have seen that there is strong evidence that large areas of the countryside over most of the Balkans had become seriously depopulated during the sixth century. Despite Slav settlement of parts of the area, the seventh-century landscape of the region still seems to have been sparsely populated". According to Tibor Živković's The Urban Landcape of Early Medieval Slavic Principalities in the Territories of the Former Praefectura Illyricum and in the Province of Dalmatia (ca. 610-950) (2013), the earliest relations between the Romans and the Slavs reveal it was not peaceful. Archaeological evidence "has shown that many large settlements became abandoned during the first half of the seventh century and that inhabitants retreated toward the remaining cities in the Dalmatian coast and its islands", and it is considered that the "influx of the Slavs should not be measured in hundreds of thousands, but rather tens of thousands in the entire vast region from Istria to Ulcinj". Sedov (1995) notes that the native population was pushed to the mountain or coastal areas and that in many regions established good relations with the Slavs, while Zupan (2013) notes that part of the "territories were abandoned by the indigenous population ... small pockets of autochthonous Romanized populations, who survived in fortified towns on higher grounds ... was either assimilated or enslaved".



In the historiography is a common theory that the Slavic migration to Southeastern Europe happened in two waves, of which the first was from the first half of the 6th century to 630, and the second migration which happened since 630s, specifically the arrival of the Croatian and Serbian tribes, however, it is also argued that Slavic migrations did not finish until the Hungarian conquest of Pannonia in early 10th century. There's a scholarly debate about the chronological order of the waves, one across Pannonia from the North and the other over the Carpathian Mountains from the East, and the "the prevalence of one over the other and the impact they had on the contemporary population" (Zupan et al. 2013).

According to Váňa (1983) "The main flow of Slavs who colonized the Balkan peninsula moved from the Ukraine along the Carpathians and across the Danube. They went as far as the Adriatic Sea, the Peloponnese, the Aegean islands and the coast of Asia Minor", while according to Sedov (1995), from the 510s began raids of Antae and Pannonian Avars to the province Scythia Minor, and soon all over provinces of Thrace, Illyria, Dacia, Moesia, Dardania, Macedonia, Dalmatia, Epirus and that the South Slavs are a mixture of both Sclaveni and Antae, but the majority is descending from the latter. 

Váňa 1983
Váňa (1983) noted that "Historians and philologists are still engaged in a lively discussion on the share of Western or Eastern Slavs in this expansion, but it seems that those arguments are well-founded according to which the main stream of migration arrived from the east across ancient Dacia in the middle of the sixth century. The starting point of a lesser wave of migration were the settlements of Croats north of the Carpathians in the upper Vistula valley, from where smaller tribal groups moved south. This movement was recorded as an old legend by the Byzantine Emperor Constantine Porphyrogenitus in the middle of the tenth century. The fact that the Southern Slav tribes bear the name of that numerically relatively small Western Slav people shows that the Croats and with them the Serbs played an important role in the struggles against the Avars; and the Slavs managed to gather around them tribal groups to which their name was applied ... The picture that archeology provides on this problem is far from complete. But it tends to support the views of historians convinced of the eastern origin of the Southern Slavs rather than the conclusion of scholars who assume a considerable influx of Western Slav tribes to the Balkans. So far no definite find has been made on the territory of Yugoslavia of the Prague type, which would indicate a movement from Central Europe. The finds of early Slav hand-made pottery are very rare in this region and tend to resemble the Romanian, which is of Eastern Slav origin".


Slavic migrations routes per toponyms by Jordan Zaimov (1967), Sedov 1995

Sedov (1995) argues that in the Middle Danube basin was a mixture of Avaro-Slav Danube type of culture and Antae Penkovka type of culture, but also some Korchak culture, and from there migrated the main wave to the South; that exist rare findings of objects and ceramics of Korchak culture on the territory of Croatia, Bosnia and Herzegovina, and Serbia which was dated to the late 6th and early 7th century, that those Early Slavs very soon incorporated Roman technologies and way of life because of which is difficult to archaeologically identify them; who do not make continuity with Slavic archaeological findings from the 8th-9th century. In conclusion, Sedov's argument suggests different scenarios, that Croats and Serbs arrived or as numerous subjects of the Pannonian Avars who later rebelled against them which is contradicting to DAI or opponents of the Pannonian Avars who encountered already settled Slavic population and assimilated them.

Straight lines are Penkovka and Ipotesti-Candesti, crossed lines Avarian, while dotted are Sukow-Dziedzice and Korchak cultures, Sedov 1995

© LIKBEZ. Історичний фронт

On the other hand, processual and post-structural & postmodern archaeologists and historians tend to overestimate the social, political, and cultural influence and underestimate population replacement. Such a revisionist viewpoint is based on ideological rather than scientific grounds and is not generally accepted (see Critical theory#Criticism and Vienna School of History#Criticism). Walter Pohl stated in Regna and Gentes: The Relationship Between Late Antique and Early Medieval Peoples and Kingdoms in the Transformation of the Roman World (2003) in regard to the Pannonian Avars that the "names were also connected with prestigious traditions that directly expressed political pretensions and programmes, and had to be endorsed by success. In the world of the steppe, where agglomerations of groups were rather fluid, it was vital to know how to deal with a newly-emergent power". Florin Curta in The Making of the Slavs: History and Archaeology of the Lower Danube Region, c.500–700 (2001) and D. Dzino in Becoming Slav, Becoming Croat. Identity Transformations in Post-Roman and Early Medieval Dalmatia (2010) and The rise and fall of the Dalmatian ‘Big-men’: Social structures in Late Antique, Post-Roman and Early Medieval Dalmatia (ca. 500-850) (2014), argue that ethnicities are not an immutable and static social phenomenon with the same core yet they are fluid, that individual findings are mistakenly extrapolated as representative of a collective ethnic group and should not be considered as equivalent through time in an attempt to prove a population replacement as a reflection of a contemporary nationalistic perspective to a distant period of time. Curta (2018) even argues that "there is no mention in any source of Slavs crossing the Carpathians, but maps of the Slavic migration have large arrows either around or across the mountain range separating northern from Central Europe, between the Upper Elbe and the Upper Dniester rivers ... To track that Transcarpathian migration, archaeologists use pottery, specifically the handmade pottery known, since the 1940s, as the Prague type", which according to An ironic smile: the Carpathian Mountains and the migration of the Slavs (2018), that there "are considerable differences between different regions adjacent to Carpathian Mountains, both in the general terms of the archaeological record, and in the specific details of ceramic morphology concerning the handmade pottery hastily attributed to the Prague type. Such differences do not justify either the use of pottery as a chronological and cultural marker, nor the use of the phrase Prague culture". Curta stated in Four questions for those who still believe in prehistoric Slavs and other fairy tales (2015) that according to archaeological data various theories do not explain the seeming lack of archaeological data that the Slavs left their Urheimat in Western Ukraine, which supposedly would reduce the data and demographics there yet it was the opposite. That the material culture attributed to Early Slavs in Slovenia and Croatia has nothing to do with that of the sites in Northwestern Hungary, Romania, and Ukraine, yet "by contrast, these early 7th-century cremation burials have very good matches ... from a number of similar sites in Southwestern Germany and Hungary". There's suspicion of radiocarbon dating results for 6th and 7th-century ceramic assemblages to the point that "migration took place at a much later date and, most likely, from a much shorter distance than commonly assumed ... from Southwestern Slovakia or, across the mountains, from Southern Poland ... something done with the approval, if not under the supervision of the Avars, perhaps in the aftermath of the civil war that shook the Khaganate in the 630s".  However, such an approach and model about a small barbarian elite culture on the frontier which through language spread Slavic ethnic identity among more numerous native population, while in the case of Curta who argues that the Slavs were invented by the Byzantines, is too simplistic and subjective narrative which is, in addition, completely contradicting the accounts written in historical sources and anthropological & genetical data. It contributed to the understanding of the complex social and cultural situation and interaction at the Roman frontier, especially for some specific regions, but does not provide a convincing general model at all.

The expansion and area of settlement by the Slavs c. 500-1000 by The Penguin Atlas of World History Vol. 1, p. 110 (25 May 2004 Penguin)

Hydrography of the Carpathian Basin was different and strongly influenced migration and inhabitation before water regulations in the 18-19th century, Arcanum


Alexis P. Vlasto in The Entry of the Slavs into Christendom: An Introduction to the Medieval History of the Slavs (1970) considered that "in course of time both Croats and Serbs became the political nuclei of larger areas in the Balkans, dissolving into the mass of Slav tribes already settled in those parts but imposing their own names", while John Van Antwerp Fine Jr. in The Early Medieval Balkans: A Critical Survey from the Sixth to the Late Twelfth Century (1991) that "the Serbs and Croats whom Constantine mentions were the second migration of different people who do not seem to have been particularly numerous. In his discussion, Constantine describes which element emerged as the ruling one over which particular territories ... and some of these Slaveni were ruled by a Croatian military aristocracy and some by Serbian one ... to dominate the disorganized Slavic tribes. They were able to provide a ruling class and be a source of unity for the different Slavic groups. Soon the newcomers came to provide a general name for all the people" but "if few Slaveni had come into the Western territories under the Avars, probably many more did so now, in the wake of the new conquerors".

In conclusion, as noticed and explained by Heather (2009), "if this much is plausible, the seventh-century Serbs and Croats were not whole peoples responsible for the complete repopulation of these parts of the Balkans. As we have seen, the better-documented instances of first-millennium migration have never thrown up a case of total demographic replacement: some indigenous population elements always survive ... It is also unclear whether their arrival represented a further major wave of Slavic immigration into the north-western Balkans, or whether they functioned essentially as an organizing element for Slavic groups already present there but formerly subject to Avar domination. If the latter, this would make them not unlike the Bulgars of the eastern Balkans ... Serbs and Croats might represent yet a third type of migrant group caught up in the Slavic diaspora of the sixth and seventh centuries ... Avar campaigns between 570 and 620 were many and varied, and this would provide a plausible context for a further bout of sociopolitical evolution among those Slavs caught up in this latest nomad war machine to establish itself in central Europe, sufficient to produce this third type of Slavic migrant group that was either large enough or militarily specialized enough to throw off Avar domination. It might be the same kind of force as the five thousand militarily ‘elite’ Slavs who launched a surprise attack on Thessalonica. If so, Serb and Croat migration might have taken the form of a kind of elite transfer, with a militarily effective force breaking out of the Avar Empire and establishing its own niche in the Balkans".

"A comparison of the historical and archaeological evidence thus sets up a seeming paradox. Those regions of Europe that saw the complete transfer of a Korchak-type material-cultural system also witnessed a Slavic migration process involving only very small social units. On the other hand, the historical evidence for much larger Slavic social units on the move (whether ‘tribal’ or military specialists – if such, indeed, were the Serbs and Croats) relates to those areas where archaeologists have not uncovered any largescale transfer of ‘complete’ Korchak-type socioeconomic systems. This is at first sight surprising. The larger the migration unit, you might suppose, the greater its capacity to import and maintain its own distinctive way of life. When you think about it, though, the larger Slavic social units were actually very recent creations, generated by processes of rapid sociopolitical and economic development that were unfolding among those Slavs closest to the Roman frontier or who became caught up in the Avar Empire ... In other words, it was precisely the larger Slavic groups rather than the small-scale farmers of the Korchak world who would have been more open to the kinds of influences and processes that would have caused their patterns of material culture to evolve away from older Korchak-type norms".

"These conflicts were underway in the later fifth and the earlier sixth century, precisely when Korchak Slavic-speakers were spreading westwards from the Carpathians and must have eased their takeover of Moravia and Bohemia. There may also have been a second political dimension to the motivations of Korchak groups. As we have seen, these migrants, moving as small-scale farming communities, need to be distinguished from the larger and more militarized Slavic entities that were simultaneously evolving further east and south through direct contact with the East Roman Empire. Given this, the Korchak-type migrants may also have been on the move so as to avoid being sucked into the orbits of these new and more powerful Slavic polities ... The militarizing Slavs of the Carpathian region thus made potentially useful subjects for the Avars, who quickly attached some of them to their train".

"On the other, Avar domination was itself something that many Slavic groups wanted to avoid – or to throw off, having once fallen foul of it. The Serbs and Croats who settled in the Balkans reportedly did precisely this, as we have seen, as did the Sorbs further west under the leadership of Samo ... Avar domination thus provided yet more reasons for Slavic groups to move out of the Carpathian and Middle Danubian regions. First, while the initial spread of Korchak-type communities clearly had other origins, having begun before the Avars became a factor, their further spread from Bohemia towards the Saale and beyond the Elbe after the mid-sixth century will have had the extra motivation of seeking to avoid absorption into the exploitative Avar Empire. This may well have prompted the spread of Slavic speakers northwards into Poland at more or less the same time. Second, Avars were responsible for the spread of the larger 'tribal' Slavic communities into the Balkans after 610, which would have been impossible if the former had not destroyed Roman frontier security. But these were the same Slavs who had been alternately fighting and serving the Avars over the previous fifty years, so there is every reason to suppose that they also wanted to put themselves, not to mention their wives and children, out of the latter’s reach".

Approximate location of Slavic tribes and others between 6th-10th century, Creative Commons
© LIKBEZ. Історичний фронт

In regard to population genetics, obviously, not all regions of Southeastern Europe had the same ratio of native and Slavic population as in the territory of Eastern South Slavs existed higher percentages of natives. Considering the majority of the Slavs came via the Eastern-Central route, the lower percentage of Slavic ancestry in contemporary populations in Bulgaria and North Macedonia does not imply that the number of the Slavs there was less numerous than among the Western South Slavic countries, on the contrary, in the medieval hinterland of the territory of Western South Slavs most probably was a state of desolation which population bottleneck produced a founder effect of a Slavic population resulting in high frequency but lack of variance in both R1a and I2 haplogroups and a higher proportion of Slavic autosomal ancestry.

3.3. South Slavic languages and dialect groups:

Besides historiography and archaeology linguistics can also tell us a lot about the origins and migrations of the Slavs. According to Zbigniew Gołąb's work The Origins of the Slavs: A Linguist's View (1992), "the territories located north of the Carpathians became in the fifth-sixth century A.D. the basis from which the great southward movement of the Slavs started, i.e., to Moravia and Bohemia, and to Pannonia and Dacia (Transylvania) ... Trubachyov has shown that in the Upper Dniestr basin there is a characteristic concentration of two layers of hydronyms: those with possible West Balkan connections (Illyrian?) and those with South Slavic connections. On the opposite side of the Carpathians (in the Upper Tisa basin) we again have a characteristic South Slavic hydronym Topla, i.e., 'warm' (river), etc.). Such facts suggest that the main route taken by the ancestors of the Southern Slavs in their migration south towards the Danube was the ancient trail through the central Carpathian passes, and the region where a kind of Proto-South Slavic linguistic comunity developed (before its split into the western, Sloveno-Croato-Serbian, and the eastern, Macedo-Bulgarian, branches) was northeastern Pannonia, i.e., later sub-Carpathian Ruthenia. From that transitional stage the Southern Slavs later moved to the Danube River along two separate routes, one more western, another more eastern (through Transylvania)". According to Jürgen Udolph's work Expansion slavischer Stämme aus namenkundlicher und bodenkundlicher Sicht (2016) on the distribution of toponyms and hydronyms the center of the ancient homeland of the Slavs was between Bukovina and Krakow, which agrees with previous conclusions. Whence the expansion of the Eastern Slavs passed by the Pripyat swamp all through central (European) Russia especially to the north and east; whence the expansion of the Western Slavs was through Bohemia to Saxony and Thuringia, and through western Poland to Brandenburg and Mecklenburg-Western Pomerania; whence the migration of the South Slavs took roughly through two main but separate waves, one western approximately through the Moravian Gate to Austria, Slovenia and Croatia to Albania and western Greece, and the other eastern along the eastern Carpathians to the Iron Gates on the Serbian-Romanian border and further to Bulgaria, Macedonia, and Greece. Along with them, there are a couple of other vaguely defined routes that encompassed the western and eastern lowlands of Hungary.



According to Kushniarevich et al. (2015), the "initial division of Proto-Slavic remains unresolved: a ternary split into West, East, and South dated to around 1900 YBP is suggested in the consensus phylogenetic tree. Further diversification of the Slavic languages took place around 1300–1500 YBP, followed by shaping of the individual languages 1000–500 YBP. Our reconstruction suggests the existence of several intermediate clades–Ukrainian/Belarusian within East Slavic, Czech/Slovak and Polish/Kashubian within West Slavic–whereas a ternary structure is suggested for Serbo-Croatian, Bulgarian and Macedonian within South Slavic. Modern Slovenian, due to its vocabulary exhibiting a significant level of a mixture with West and South Slavic languages, was excluded from the lexicostatistical analysis".

Kushniarevich et al. 2015

According to Ranko Matasović's work Poredbenopovijesna gramatika hrvatskoga jezika (2008), the "area on which was spoken the language from which developed Proto-Slavic before the expansion to the Danube limes of the Roman Empire most probably was located Northeastern of the Carpathian Mountains; there was the territory with the greatest density of hydronyms which can be related to Proto-Slavic, and exactly from that starting point it is easiest to explain later Slavic migrations ... that area between rivers the Dniester in the South, Western Bug in the West and Pripyat Marshes in the North, matches well enough with widespread archaeological culture Korchak from 5th to 7th century. It later in the 7th-century blends into Prague or Prague-Penkovka culture, which is without doubt Slavic, attested in the area of Slavic expansion to the West, specifically Bohemia and Moravia". According to Matasović (2008), the historical reality is, or should be, reflected in the genetic relationship between languages in linguistics. Regardless of doubts, for the phylogenetic tree of Slavic languages (see below) it is considered that existed Proto East Slavic-South Slavic language, while it is not clear whether existed Proto-West Slavic with common innovations for all and only West Slavic languages. The first branched into the South Slavic and East Slavic language families. A variation of Proto-Slavic language was spoken since the time of arrival of Slavs in the 6th and early 7th century until the 9th-century when began a diversification. However, there still exist some doubts about the existence and uniformity of the Proto-South-Slavic language, and if did it probably did not last long because there was no known cultural and political unity of the South Slavic area nor it is bounded by natural boundaries. According to Łowmiański (1964) and Majorov (2006), South Slavic is also more related to the Eastern Slavic language group which indicates the starting point of the migration towards the Balkan peninsula.

Matasović (2008)

According to Mijo Lončarić's work Prilog južnoslavensko-zapadnoslavenskim odnosima, s posebnim obzirom na slovačko-kajkavske veze (1993), from a linguistical point of view is considered that the 6th century was a turning point of innovation and development of the Proto-South-Slavic language. From then until the 9th century existed a Slavic dialect continuum in Central and Eastern Europe; West Slavic and East Slavic had their continuum in the north while in the south both West Slavic and East Slavic had a continuum with South Slavic language. The triad boundary was probably located in the territory of central Slovakia. Proto-Slavic language and sub-groups most probably were not unique & the same on the whole territory yet like all-natural languages had local dialectological differences.

Approximate early medieval prevalence of Slavic languages subgroups continuum and contact zones

Slavists like Fran Ramovš since the 1930s argued that the Slavs "may have arrived in the Carpathian Basin with some preexisting dialect divisions" (Grant H. Lundberg 2013), that the founder population of the specific dialectological-geographical groups spoke a variety that was structurally different from neighboring Slavic speakers, although eventually would make a mutual continuum, and can point in some detail the routes of the founder populations. For example, some linguistic innovations connect pockets of Prekmurje Slovene, central Bulgaria, and central Russia (Avgust Pavel 1942 & 2020); Slovenian and some Croatian with Czech and Slovak, Slovak with Croatian and Slovenian, Czech with Slovenian, Middle Slovak with some Croatian (Chakavian and Western Shtokavian), Middle Slovak with some Croatian (Kajkavian) and so on (Lončarić 1993).

According to many linguists including Ramovš, Pavle Ivić as well as Mijo Lončarić's work Rani razvitak kajkavštine (1988), from the 6th until 9th-century semantic isoglosses show that on the level of language groups, languages, dialect groups, and dialects happened: first the division of the South Slavic language group into the Southwestern and Southeastern subgroups. From the former will later develop Serbo-Croatian and Slovenian while from the latter Bulgarian and Macedonian languages. While the latter is arguably very uniform with the main difference being between Western and Eastern Bulgarian (with Western used as a base for Macedonian), the former is diverse with at least two up to five proto-dialectological-territorial divisions. According to isoglosses, the Southwestern subgroup is secondly divided into two subgroups, Western-Southwestern Slavic and Eastern-Southwestern Slavic. From the Western-Southwestern Slavic will emerge four independent subbranches (Slovenian, Kajkavian, Chakavian, and Western Shtokavian dialect groups) while from Eastern-Southwestern Slavic will emerge one or two independent subbranches ("true" Shtokavian i.e. Eastern Shtokavian and transitional Torlakian dialect groups). However, the Eastern-Southwestern Slavic dialects also share an old innovative isogloss with Bulgarian which archaic form is preserved by other Western-Southwestern Slavic dialects. By adding another isogloss the division of Western-Southwestern Slavic wouldn't be in two but three subgroups; Western-Southwestern Slavic (Northwestern part of Slovenian language), Central-Southwestern Slavic (Southeastern part of Slovenian language, Kajkavian, Chakavian, and Western Shtokavian dialect group), and Eastern-Southwestern Slavic (Eastern Shtokavian and Torlakian dialect group). The branching of Western-Southwestern Slavic can be phonologically additionally argued that chronologically first separated Slovenian, soon Kajkavian and lastly Chakavian and Western Shtokavian (Iva Lukežić 2012). By the 9th or 10th century were formed proto-units of these dialect groups while in the form as are known today only since 12th and later centuries which complicates historical-genetic argumentation.


Lončarić (1988)


As said by Dalibor Brozović and others like Marko Filipović in South Slavic languages with Neo-Shtokavian base (2014), in conclusion, the Proto-Southwestern Slavic (ZJP) language shouldn't be seen as a single entity, but rather composed of four proto-dialectological groups which got their names on geographical location upon the arrival to the Balkans, and that is Alpine (Al) represented by Slovenian dialects, Pannonian (Pa) represented by Kajkavian dialects, Littoral (Pr) represented by Chakavian dialects, and Dinaric-Rascian (Dr) represented by Shtokavian and Torlakian dialects - more correctly into five groups, Alpine would be Slovenian, Pannonian would be partly Kajkavian partly Western Shtokavian, Littoral would be Chakavian, Dinaric would be minor Chakavian with major Western Shtokavian, minor Dinaric-major Rascian would be Eastern Shtokavian and Torlakian. In addition, could be added a sixth Balkan group composed of Proto-Southeastern Slavic (IJP; Bulgarian and Macedonian dialects).

Filipović (2014)

Since the early times, the Chakavian dialect group was never completely unified and possibly never existed, being only a collection of archaic dialectological remains when excluded Kajkavian and Western Shtokavian dialect isoglosses, with the North Chakavian dialect having a tendency toward Kajkavian and Slovenian while Middle and Southern Chakavian toward Schakavian i.e. Western Shtokavian; Shtokavian dialect group from the early times was disunited with Western Shtokavian being closer to Chakavian and Kajkavian while Eastern Shtokavian (and Torlakian) with some old phonetic isoglosses having a continuum with the Bulgarian language (Ivić 1957-58, Lončarić 1988, Vergunova 1996). In the development of today's Shtokavian dialect group happened a rare case of the development of proto-units where development does not flow divergently, but convergently - differences do not increase but decrease (this was due to historical and political conditions especially since the 16th century).

Croatia in 10th century by Croatian Encyclopedia; boundaries of phonological isoglosses by Ivić 1958; premigratory and postmigratory distribution of Serbo-Croatian dialects by Brozović 1988/98 


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As was stated, the chronology of semantic isoglosses and as such changes which began and finished between the 6th and 9th centuries were argued or to have begun in the Slavic homeland and finished in the Balkans or to have begun and finished in the Balkans (Lončarić 1988). The lexical isogloss division also depends on distance but it is thought to be an open system that is diffuse without strong isoglosses i.e borders with different overlaps between languages (for e.g. South Slavic with East Slavic, South Slavic with West Slavic, Serbo-Croatian with Bulgarian, or Serbo-Croatian and Slovenian, Southeastern Slovenian and Southwestern Kajkavian and Northwestern Chakavian lexicon and so on). However, when such borders do exist can be drawn on the basis of statistical analysis and frequency of differences in the lexicon. Until recently did not exist detailed lexical research to give a basis and confirm the previous proto-territorial-dialectological units division by phonetical isoglosses (Lončarić 1988). Nikita Ilyich Tolstoy was among the first linguists when in 1962-1976 on the basis of lexicon reconstructed historical division of Serbo-Croatian dialect continuum and showed opposition of central and lateral South Slavic zone indicating settling of the Balkans by at least two different routes. He also researched Chakavian-East Slavic comparisons and found in geographically distant Slavic dialects evidence of old Proto-Slavic dialectological connections. Many others followed in his steps, with Oleg Trubachyov concluding that linguogeographic research of Slavic lexicon can reveal by lexical differentiation very old Slavic dialectological connections and relations (Yakushkina 2020).

Ekaterina Yakushkina in her research Isoglosses of lexemes of Proto-Slavic origin in the Serbian-Croatian areal (2020) found that by comparison of many Proto-Slavic lexemes, from eleven basic thematic groups, the Serbo-Croatian dialects are roughly divided into eight zones in which are opposed by lexical isoglosses:

1. Most of the Shtokavian dialects / Kajkavian / Chakavian and part of the Zeta-Raška Shtokavian dialect of the so-called "side belt" (1 isogloss).
2. Kajkavian and Northern Chakavian / Shtokavian and Middle Chakavian and Southern Chakavian dialect (3 isoglosses).
3. "Side belt" (Chakavian and Zeta-Raška dialect) / most of the Serbo-Croatian dialects (3 isoglosses).
4. "Side belt" and Kajkavian dialects / most of the Shtokavian dialects (2 isoglosses).
5. Kajkavian and Chakavian dialect group / most of the Shtokavian dialects (2 isoglosses).
6. Western dialects (Kajkavian, Chakavian, Western Shtokavian) / Eastern dialects (Eastern Shtokavian, Torlakian) of Serbo-Croatian continuum (8 isoglosses).
7. Dialects of Serbia / other Serbo-Croatian dialects (1 isogloss).

It leads to the conclusion that "most of the lexical differences, probably actual for Proto-Slavic, contrast the Western and Eastern parts of the Serbo-Croatian territory (*kruxъ –*хlěbъ 'bread', *оbtokъ – *obstrovo 'island', etc.), and usually, the source of lexical differentiation is an innovation that spread from the West. Other types of areal oppositions of Proto-Slavic lexemes on Serbo-Croatian territory are connected with the opposition of Shtokavian dialects to other dialects. The proto-Slavic lexical isoglosses also share Chakavian and Kajkavian dialects (sometimes North Chakavian are combined with Kajkavian), the main part of Shtokavian dialects and South-Montenegro dialects". It implies that lexical isoglosses mostly confirm proto-territorial-dialectological units division of phonetical isoglosses especially the division of Southwestern Slavic into Western-Southwestern and Eastern-Southwestern particularly within the Serbo-Croatian continuum.

Ludmila Vergunova in her dissertation The geographic distribution of Proto-Slavic dialectisms and the genesis of the South Slavic languages (1996), also citing Ivić, Lončarić, and others argue that "traces of the ancient Proto-Slavic dialects are evident on the level of smaller dialectal units and regional vernaculars. The isoglosses (mainly lexical and semantic), connect modern South Slavic subdialects with different territories of the northern Slavic world" and in her goal to find additional arguments to support a hypothesis that rejects the idea of a homogeneous Proto-Slavic or the direct development of modern Slavic dialects from Proto-Slavic ones came to the conclusion that "the Slavic colonization of the Balkan peninsula was actually a starting point for leveling tendencies which with time brought the rise of a Balkan linguistic unity and not the beginning of a dialectal division of South Slavic dialects". 

Among the detailed findings was a sharp division of Carpathian dialects into Western and Eastern Carpathian. The western part roughly includes West Slovak dialects while the eastern part Ukrainian Carpathian and sub-Carpathian dialects and East Slovak dialects. It implies that the traditional three semantic divisions of Slovak dialects are contrasted by lexical and ancient dichotomous division of the Slovak territory into the north-east and south-west, while the dialects of the latter region differ sharply from Ukrainian dialects east of the Dniester river and Polesia. This region has numerous correspondences with the Southeastern part of the central Balkan peninsula. North-eastern Slavic dialects (Northwestern Ukrainian, Western Belarusian, Pskov, and Novgorod) reinforced the previously observed connections between the Northwestern Russian territory (Pskov-Novgorod land) and the West Slavic and Slovenian languages. The modern dialects of the assimilated Slavic languages retain traces of the Proto-Slavic dialect division (for example numerous borrowings in Romanian and Hungarian dialects).

As if it were a projection of the Carpathians, the central South Slavic area made by Serbo-Croatian dialects also splits into two parts, Western and Eastern-Southwestern Slavic. The East Carpathian correspondences are characteristic primarily for the Eastern but also the Western part and from there to the periphery. The central South Slavic area is opposed to the so-called Balkan lateral area (West Slovenian, insular Chakavian, and partly Macedonian dialects) who do not share ancient lexical and semantic isoglosses with the East Carpathian region. Within the central South Slavic area the Schakavian (Western Shtokavian) dialects appear in the borderline role between Western and Eastern-Southwestern parts, while Kajkavian and East Slovenian dialects are characterized by the absence of many East Carpathian Slavisms and old Balkanism substratum. It seems that East Slovenian and Kajkavian dialects have not experienced the strong influence of the Romanized or non-Romanized Daco-Thracian shepherd population, which is to some extent related to the geography of the spread of these dialects (the territory of the Romanized Illyrian tribes). It possibly indicates a later or different starting point of the Slavic route migration of Proto-Kajkavian speakers.

Vergunova 1996

PCA based on Eurogenes K13 results of hundreds of nationally and regionally confirmed genetic genealogical samples. The proximities show significantly lower Roman-Balkan substratum among Kajkavian Croats and Slovenes compared to other South Slavic regional populations

E. I. Yakushkina in About Lexical Differences between Old Shtokavian Croatian and New Shtokavian Serbian Dialects (2020) confirms that "among the lexical differences there are a number of Proto-Slavic synonyms, which presumably contrasted various proto-Serbo-Croatian dialects at the beginning of the second Millennium ... from geolingustic point of view Slavonian and Vojvodina dialects present an opposition of Western and Eastern Shtokavian". Marinka Šimić in Lexical archaisms in some Western Shtokavian dialects (2021) also confirmed that "the Western Shtokavian Ikavian dialects have a common Chakavian-Shtokavian stratum" and "conclusion that preserve a significant layer of lexical archaisms, similar to the Chakavian and Kajkavian ones, which indicates their common origin".

Pavao Krmpotić in Prilog istraživanju naslijeđenoga slavenskog leksika u hrvatskim kajkavskim govorima (2015) found some Kajkavian lexemes were shared in old Chakavian and some Shtokavian dialect groups, some in West and East Slavic and Slovenian language, some in West Slavic and Slovenian language among others. In conclusion, the "existence of such examples is a piece of evidence that the Kajkavian and Slovenian dialects made a dialect continuum with West Slavic languages before the arrival of the Hungarians to Pannonia".

András Zoltán in his Славянские диалекты Карпатского бассейна во время прихода венгров (IX в.) (2013) investigated the oldest Slavic borrowings to Hungarian with regard to the reconstruction of the dialectal landscape of Slavic in the Carpathian Basin at the time when the Hungarian tribes arrived here at the end of the 9th century. According to some phonetic criteria, it seems that the Hungarians found two main Slavic dialects in their new homeland: the Pannonian Slavic with mixed West and South Slavic features and the Bulgarian Slavic. It shows that in the area of the Carpathian Basin there was a linguistical continuum at the macro level of West and South Slavic and at the micro-level of South Slavic dialects. 

Mijo Lončarić in The Borders of the Kajkavian dialect in the past (1995) wrote that "before the Hungarians came to Pannonia there existed an uninterrupted language continuum between [West Slavic and South Slavic] language groups ... before 10th-century one cannot speak about dialectal affiliation, it is questionable whether one can speak about language affiliation at all. Namely, the Slovenian language was not at the time very different from other parts of Western South Slavic Proto-language. It is certain that across the Drava river on its left river bank in today's Hungary the language couldn't be different, at least not considerably different, from that on the right river bank. One can certainly determine that the language of the Slavs in today's Hungary belonged to the South Slavic language group. The middle Slovakian dialect is in its basis also South Slavic for it shows many important South Slavic features. After the Slavic language continuum had been broken in Pannonia and after the separation of Middle Slovakian speeches from their South Slavic motherland these speeches were under the influence of the West Slavic majority and they developed further in that direction". Lončarić concluded that North of the Lake Balaton expanded the South Slavic language and that if its unity was not interrupted there would have developed a new type of Proto-Kajkavian dialect. 

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In addition, as stated by Rafaela Šejić in Mjesto akcenta u ruskom i čakavskom (čakavština srednjodalmatinskih otoka) (1999) and many other linguists, the Chakavian is the most archaic Southwestern Slavic dialect group and it is needed its knowledge with Old Church Slavonic for the phonetic research of problems in Proto Slavic. It preserved traces of deep Slavic antiquities in all its richness and hence is especially significant for Croatian and Slavic studies. It indicates that the Croatian and most Bosnian and Herzegovinian speakers of Chakavian and Western Shtokavian came as mostly common folk of mainly East Slavic ancestry with a slight West Slavic continuum; that the White Croats and probably Lendians in Zahumlje among others did not come during the second wave yet first with the Pannonian Avars and as stated in DAI successfully fought against them because of which had to be strong enough tribes for such an act; kind-of non-Slavic personal names of Croatian mythological five brothers and two sisters which "led" the Croats and of the first Croatian ruler Porga also confirm Croatian-Avar first wave contacts which shows that DAI's consideration about the 7th century arrival of the Croats was based on author's knowing of their 9th and 10th location in Eastern and Central Europe; confirms the claim from DAI that part of the Croats set off and expanded power to Pannonia only after their arrival to Dalmatia implying some difference between Slavs settled in early medieval Croatia and Bosnia (Chakavian-Western Shtokavian) from those in Lower Pannonia (Kajkavian); confirms legend about the seven or eight tribes who arrived roughly from Poland to Croatia and Zahumlje; confirms close political connections between early medieval Croatia, Pagania and Zahumlje, putting of the "Serbo-Croatian" border on river Neretva, mention of Croatian ethnonym up to Duklja, political hostility between Zahumlje and early medieval Serbia; that as stated in some historical sources and by many historians early medieval Bosnia was not part of early medieval Serbia neither population of Bosnia, Pagania, Zahumlje etc. was of Serbian ethnic origin; early tight connections betwen Croatia and Bosnia and so on. Polabian tribes of the Serbs and Abodrites were the ones that most probably came during the second wave because of which in DAI isn't stated to have fought the Avars but rather settling in areas previously desolated due to the Avars, escaping rather Frankish pressure during Samo's rule and effects of Late Antique Little Ice Age. Serbian tribe partly settled in the territory of what would be Eastern Shtokavian dialect group but didn't bring such dialect because the territory was mainly inhabited by other Slavic tribes and was initially controlled by the Bulgarians neither today's speakers are mainly descending from the Serbian tribe for the reasons stated in previous chapters above, having an old connection with former and current Bulgarian Slavs of East Slavic continuum. Perhaps the West Slavic isoglosses in some Serbian and Macedonian dialects could be preserved remnants of traces of the Serbian tribe from Polabia (having analogies in DAI where's stated that when they arrived at the Balkans settled first an area near Thessaloniki, also the 7th-century formation of the Asia Minor city or region of Gordoservon which was inhabited by Macedonian Slavs).

Interestingly enough, according to Zupan (2013) "the differentiation of the Slovenian population from the rest of the Balkan populations is based primarily on the lower frequency of the I2a1b-M423 haplogroup. The calculated age of this specific haplogroup together with the variation peak detected in the suggested Slavic homeland could represent a signal of Slavic migration arising from medieval Slavic expansions. However, the strong genetic barrier around the area of Bosnia and Herzegovina, associated with the high frequency of the I2a1b-M423 haplogroup ... a medieval Slavic signal from modern-day Ukraine ... Furthermore, the genetic proximity of the Slovenian, Slovakian and, to some extent, the Czech populations, whose territory was united in the early days of Slavic settlement under the Samo’s tribal union, is of particular note (Barford, 2001). The homogeneous genetic strata of these three Slavic countries could reflect a common ancestral Slavic population in the central European region, which was later divided by the settlement of German tribes in the region of modern-day Austria and was influenced (mostly in the Slovenian population) by the South Slavic genetic pool ... AMOVA revealed the highest variation between groups when the Slovenian population was grouped together with other West Slavic populations, suggesting that the West Slavic populations and the Slovenian population are more genetically similar to each other compared with the South Slavic language group, which consists of the Slavic Balkan population and Slovenian population. This finding favors a hypothesis that suggests an origin of the Slovenian language in the West Slavic language group (Bezlaj, 1967; Grafenauer, 1950; Žužek, 2007)"

The Slovenians who are genetically very close to West Slavs have a significantly lower frequency of M423 (~13%) and especially I-PH908 which separates them from other South Slavs and as such the Western route of migration for the main percentage of I-PH908 among South Slavs is historically and linguistically highly improbable. It also raises the question of whether the Pannonian linguistic population is somehow related to the Dulebes who lived between Lake Balaton and the Mur River in today's Hungary, and late 9th and early 10th Moravian migrants to the Pannonian part of Croatia, and to what extent were genetically unrelated to other dialectological populations? Since both Slovenian and Kajkavian speakers are still mainly R1a-M558 but with slightly or significantly lower frequency of I-Y3120 and its I-PH908 subclade it implies that most probably arrived somewhere from Western Carpathians where and when the early West Slavic population was similar to the Eastern Carpathian population, and possibly happened at the time or later some additional mixing with Eastern Carpathian movement of the Slavs. In short, there were roughly three main and separate migration routes: one from Western Carpathians and two from Eastern Carpathians, one from the western and the other from the eastern part of the Carpathians. What may additionally indicate two separate Carpathian routes is that seemingly I-S17250 subclades like I-Y5596, I-Y4882, and clade I-Y4460 which are also found in the Rusyns are prevalent in the western part of the Western Balkans while the clade I-Z17855 is prevalent in the eastern part of the Western Balkans and the Eastern Balkans. A similar thing is seemingly evident with R1a subclades with some being more prevalent in Western or Eastern Balkans including transitional zones.

A map of the main migrations of the Slavs in the 6th and 7th centuries based on archaeological and linguistic data. It is also confirmed by historiographical, anthropological, and genetic data. For a high-quality map see Scribble Maps.

On the general basis of the genetic relationship and continuum between languages and dialect groups as well as the early medieval date of their formation and diversification - can be concluded that the majority of the South Slavs arrived in Southeastern Europe from the Carpathians; more from two Eastern Carpathian routes through Wallachian Plain and Pannonian Basin rather than the Western Carpathian route via Moravian Gate; the Serbian medieval tribes from Polabia should have spoken the West Slavic language but as contemporary Serbo-Croatian is not West Slavic language nor has significant West Slavic influence as Slovenian these tribes did not leave enough linguistic trace and were assimilated by the majority while the Croatian and related tribes from South Poland and Western Ukraine were perhaps only partial speakers of West Slavic but also East Slavic and left a lot of linguistic trace; the premigratory proto-units of Serbo-Croatian dialect groups partly don't correspond with modern dialect group, language, ethnic and political definitions and borders and as such cannot be used for a simplistic and nationalistic argumentation of I-PH908 migration with an individual tribe or from Central Europe and related convoluted theorizations; the relationship of the languages support the ethnogenetic discontinuity between contemporary Leipzig & Sorbian and Serbian populations.

5. Conclusion:

The results from various fields, including research of autosomal DNA, show that Slavic migration was significantly large to physically replace, depending on the Southeastern European country, between 30-65% of the old-Balkan population. The Y-DNA haplogroup I-CTS10228 > I-Y3120 > I-S17250 > I-PH908 due to its young age and distribution is particularly taken into consideration. The most likely location of origination is Central Europe and from there moved to Western Ukraine. In the analysis arises potential cultural-tribal pattern Zarubintsy culture > Kyiv culture > Kolochin, Korchak, Penkovka, and Ipotesti–Candesti cultures > of partly Antae partly Sclaveni expansion towards Southeastern and Central Europe between early 6th and mid-7th century.

The autosomal DNA, Y-DNA haplogroup R1a and I2 subclade frequency and variance, archaeological and linguistical data indicate an Eastern rather than the Western route of the main wave of South Slavic migration, specifically in regard to carriers of I-PH908 subclades. Such a conclusion is also supported by a comparison of Y-DNA haplogroup frequency and variance in contemporary Serbian and Sorbian/Leipzig population, contemporary Croatian and Western Ukrainian and Rusyn population, contemporary Slovenians and South and West Slavic populations, and particularly by the genetic relationship between South Slavic with East Slavic and West Slavic languages. It complies with Váňa's (1983) archaeological thesis that "moved from Ukraine ... the key area is the territory of present-day Romania, ancient Dacia, for the main stream of Slav colonists moved along the curve of the Carpathians and along the Danube". 

There were roughly three main and separate migration routes - one from Western Carpathians and two from Eastern Carpathians. From the Western Carpathians via Moravian Gate came the core of the South Slavs in the Eastern Alps and Western Pannonia, from the western part of Eastern Carpathians came the core of the Slavs settled in the Littoral and Dinaric zone (partly also West and East Pannonia), and from the eastern part of Eastern Carpathians came the core of partly Western and mainly Eastern South Slavs. To the former belonged so-called Seven Slavic tribes, including the Severians and many other tribes who became part of the First Bulgarian Empire and later part of the Principality, Grand Principality and Kingdom of Serbia as well those who settled in Albania and Greece. Besides that, the issue with the arrival of the Serbian and Abodrite tribes in the 7th-century second wave is that according to it was much further West of the West Carpathian Mountains where is a contemporary lack of frequency and variance of subclade I-PH908 or haplogroup I2 in general. They were assimilated by other South Slavs and left few traces in a similar fashion of a Hungarian and Bulgarian ruling elite. While the Serbian tribe came by Western route from Saxony and Bohemia, the tribes which settled in Croatia and Bosnia & Herzegovina most probably came from the western part of Eastern Carpathians and were among the first-wave South Slavs who successfully revolted against the Pannonian Avars. During the arrival and over the following centuries the contacts between South Slavs only expanded resulting in transitional and dialect continuum zones.

However, archaeological data of the South Slavic migration, or even their lack of it, has an ongoing debate and various interpretations. Extreme lack of R1a and I2 subclades variance, especially in the Western Balkans even if is taken into account as a precaution the post-migration founder effect, does not significantly contradict the post-structuralist perspective that there was no mass-migration, but in fact, shows a homogeneous and large enough population which settled on vastly depopulated territories where were encountered by a native population which already experienced a drop in population size because of several historical and environmental factors. Up till the late modern period, there was a generally low population size and density in the Western Balkans which was induced by plagues, wars, underdeveloped economies, and local migrations which in turn by founder effects caused the modern high frequency of I2 in the same region. Due to various processes, including the fact the new Slavic identity earned prestige and its elite managed to claim a political rule over independent principalities (later kingdoms), the newcomer male population also had an advantage over native males. In addition, among all South Slavic populations the matrix-nationalities, as well as languages, mostly received the ethnic identity of Hungarians, Turkic-speaking Bulgars, and Slavic-speaking Croats and Serbs, in the former case only minority groups while in latter case possibly significantly numerous groups that were imposed as a ruling class with state-building skills.