October 11, 2019

PH908 subclade in scientific studies

In the previous post, "PH908 subclades migration", some subclades are based on only two samples, but in the case of I-Y84307 both samples are from the same country and as such it cannot be presumed its movement direction to Southeastern Europe. In that regard, this post is an attempt to find the I-PH908 subclade 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 (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 (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), 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 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 as 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 analysis, the most recent common ancestor of the two YFull samples is dated to 1300, as such the MRCA of these haplotypes lived 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 (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, 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 seven SNPs currently defining the subclade. 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 post, "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 post will have a different approach. Considering the conclusion that the haplogroup I-PH908 arrived in the Early Middle Ages from the North, it will be a synthesis of the countries of origin of the I-PH908 subclades found at the YFull YTree v8.05.00 and Y-Haplogroup I2a Project (including a post from 13 March 2020) 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 NGS test and does not necessarily reflect reality.

DYS561=15:

0) I-PH908*: North Macedonia, "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", "Montenegro", "Serbia", Slovakia, "Ukraine", USA
** Y6651/Y6652: Czech Republic
** Y45843: Bosnia and Herzegovina, Croatia
** BY169115: Italy, Serbia
** A8740/A8741*: Germany, "Poland"
*** Y59865: United States

Z16983
2) A5913/A5914*: Croatia, 2x Central or Eastern Europe
* A22312*: Montenegro
** BY153567
*** BY154460/BY152858: Bosnia and Herzegovina, Montenegro
* 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*6+SNPs: Croatia

Y84307
5) FT168731*7+SNPs: Tatarstan

6) BY198275: Poland
* P41.2/M359.2
BY198275

DYS561=16

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

1) FT16449*: "Bosnia and Herzegovina", Bulgaria, Russian Federation
* Y126296: Bosnia and Herzegovina, Ukraine
** Y81557: Serbia
*** BY57773: Serbia
**** Y189253: Italy (Albania)
**** Y103938: Bosnia and Herzegovina
***** Y99196: Croatia
* FT36524: Belarus, Russian Federation
* MF2888: Croatia, Germany, Montenegro
* BY173300: Bosnia and Herzegovina
* FT138628: Bulgaria, Serbia
* Y99608: Germany, Poland
* Y151633
** FT33812: United States (Croatia), Croatia
** BY169079: Bosnia and Herzegovina
*** Y144303
**** Y144305: Austria, Croatia
**** A22770/BY170155: Croatia

FT16449
2) BY97555/Y51673*3+SNPs: Bosnia and Herzegovina, Slovakia

Y51673
3) Y57291*: Serbia

4) FT14506*: Hungary, Bosnia and Herzegovina
A13912*: Bulgaria
** A13907*/Y30729*/BY37511*: Croatia, "Hungary"
*** BY106777: Croatia
* FT177529: Poland
** FT176058: Russian Federation
*** FT260228: Poland (Galicia)
**** FT259981: Poland
* BY135769/BY14506: Poland (Galicia), Russian Federation
* FT41224
** Y52621: Bosnia and Herzegovina, Moldova
*** FT190799: Montenegro, Serbia
** BY191170/BY191770: Croatia
*** BY189804: Bulgaria
* Y56203*: Poland (Galicia)
** FT25907: Bosnia and Herzegovina
** Y134603/Y134578: Montenegro, Serbia

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. The 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 limited Polabian Slavic contribution (only for subclades 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 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
* The post will be updated every couple of months with the development of the Y-Tree (14.5.2020.).

April 10, 2019

South Slavic origin and I2a-Dinaric South (PH908)

In this post 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 linguistics, historiography, archaeology, 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 > PH908 in the male population. At the end is a summary of the conclusions.

1. Autosomal DNA:

In this part will be cited and/or 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), and shared an additional 2019 independent analysis of recent scientific 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 50% up to 60% shared ancestry with Northern populations predominantly of Slavic origin, roughly said medieval migratory ancestry, and at least 30% 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 60% shared ancestry with Northern populations predominantly of Slavic origin, with a south-north gradation, roughly said medieval migratory ancestry, and at least 30% up to 60% shared ancestry with Southern populations, with a north-south gradation, roughly said ancient autochthonous ancestry.

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

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 upcoming gene pool, and depending on the Southeastern European region the medieval Slavic component over time replaced approximately 30-60% of the native population. Throughout history, there existed theories which ranged from autochthonous to Pan-Slavic, but both were correct and wrong as overestimated or underestimated specific components in the ethnogenesis arguing for physical or socio-cultural influence.


2. Y-DNA:

According to Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe by Battaglia et al. 2009, Hungary (53) belonged to I1 3.8%, to I2a1 14.1%, to R1a 56.6% and to R1b 15.1%, which together make over 80%. However, according to Šehović et al. A glance of genetic relations in the Balkan populations utilizing network analysis based on in silico assigned Y-DNA haplogroups (2018) which analyzed 100 samples from Purps et al. (2014), belonged to I1 13%, to I2a1 9.5%, to R1a 21.2% and to R1b 21.5%, which together make over 65%, indicating one or the other has a sampling error.

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), it 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 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 is almost exclusively represented by Z280>CTS1211 subclades among Hungarians and Yugoslavs (for e.g. >YP3992 and >Y2608) with possible expansion from Poland and Czechia-Slovakia, while equally both Z280 and M458 subclades among Bulgarians and Romanians, which TMRCA and distribution "appears to be of Slavic origin". The second, also known as I2a-Dinaric South (more details in the next paragraph), is mainly represented by CTS10228>Y3120>S17250>PH908 subclades which had a very rapid formation and expansion since 1800-1500 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
Image from "Haplogroup I2" by Eupedia

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 forming at least 30-70% with south-north gradation, with some additional Celtic-Gothic or late medieval remains of R1b and I1 subbranches, it results with a very positive or almost identical correlation to the autosomal 30-60% 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 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 post from 7 November 2016). According to YFull YTree, the SNP was formed around 1800 YBP (Cl 95% 2100-1550 YBP) and had time to most recent common ancestor around 1700-1500 YBP when emerged numerous subclades (~22), many still not defined, indicating a rapid expansion in the Early Middle Ages. For now, they are divided by STR DYS561=15/16 marker results making two large sub-branches, and each has 4 immediate known subclades (see post from 4 June 2019). According to Cullen (see 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".

Its subclades represent the majority of I2 percentage found in Southeastern Europe, which is evidence that disproves the old scientific thesis that I2 is of autochthonous origin. While Dinaric South cluster is prevalent among Yugoslavs, the Dinaric North subclades are also significantly present among Bulgarians. PH908 has regional peak frequencies in Dalmatia and Herzegovina with 50-60% which indicate local founder effects, and 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.

I-PH908 Draft Tree (04-06-2019) 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 contemporary PH908 (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

Other visual attempts of contemporary and medieval placement of 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:

The paternal haplogroup I2a1 is argued to have been "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 located in Eastern Europe and 3600-2800 BCE located in Central-Eastern Europe. However, until now in ancient samples were only found I2a1 or I2a2 subclades which are not of I2a1b2-L621 lineage which "formed 11200 YBP, TMRCA 6500 YBP" (YFull YTree). It was found in 5 or 6 samples from Hungarian conqueror period (895-mid 10th century) which were positive to L621, but negative to S17250 (Neparáczki et al. 2019), the Y3120 "formed 3800 YBP, TMRCA 2200 YBP" (YFull YTree) was found in 1 sample according to preliminary DNA results from Early Medieval Poland (900-1200 CE), and 1 sample from archaeological site Sungir from Russian High Middle Ages (1100-1200) belonged to Y3120>S17250>Y5596>Z16971>Y5595>A16681 which formed circa 1700-1500 YBP (YFull YTree). 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".

The approximate date of formation, 200 CE ± 300, per se does not give a clear answer whether it emerged in Early Slavic or another ethnolinguistic group of people which would be several centuries later assimilated by then known Slavic tribes. Most probably it will be never known the tribe of origination, while the archaeological culture in the wide region also had a heterogeneous and complex history. Considering previous information and maps which placed it around Western Ukraine, there exist two options:

1) Przeworsk-Zarubintsy cultures, particularly Zarubintsy in the East, which is considered to be related to Early Slavs, but it was multiethnic with Celtic, Germanic, Scytho-Sarmatian and Dacian influences. The latter will be replaced by Chernyakhov culture, also multiethnic culture, which will be replaced by late 5th and mid-6th-century Slavic cultures of Ipotesti–Candesti, Penkovka, and Korchak.

Wikimedia Commons
2) Lipitsa culture of Dacian tribes primarily of Costoboci, who forcibly moved near tribes of Carpi and 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 expansion 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 localization it becomes a bit difficult to argue from such a population emerged a rapid, numerous, and widespread expansion which will be vital to the following Slavic culture further to the Northwest and Northeast. Perhaps it could be possible if the descendants of the common ancestor regularly changed the ethnic and social identity and managed to successfully use new political movements for their own reproduction, being swapped into Hunnic-Gothic and then Slavic or directly into Slavic expansion. Interestingly the PH908's formation and expansion happened around the time of influence of Antonine (165-180 CE) and Cyprian (249-262) and then Justinian (541-542) and Roman (590) plague, and Late Antique Little Ice Age (536-660), which positively and negatively influenced both the "barbaric" tribes and Roman Empire's population and their mutual relations.

The approximate date of PH908's numerous subclades formation, 300 CE ± 200, seemingly coincides with the Slavic movement. As previously said, most probably it will be never known the tribe of origination, but it could be presumed the 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 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 (Middle-East Slavs), and Antes (East Slavs). Considering the contemporary wide distribution of 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
According to Florin Curta's An ironic smile: the Carpathian Mountains and the migration of the Slavs (2018), the "earliest assemblage associated with the Prague culture –  the second half of the 5th century" which also included wheel-made pottery was found in the Outer Eastern Carpathians, specifically Bukovina region of Western Ukraine. However, the "three recent excavations confirm that Western and Central Slovakia were occupied through the second half of the 6th century by a population that had nothing to do with the Prague culture", and that "similar assemblages with handmade pottery appear inside the Carpathian Arc ... dated to the 6th century and have been attributed to the Gepids ... also mentioned among finds from settlement and cemetery sites discovered in Hungary and dated to the Early Avar age (ca. 570 to ca. 630)". Same thing about ceramic assemblages in Lesser Poland (625-635) and Lower Silesia (677), which concludes that the "ceramic assemblages from East Central Europe that have been attributed to the Prague culture cannot be dated before 600, either in Slovakia or in northwestern Romania. Inside the Carpathian Mountain Arc, the earliest assemblages that can be attributed to the Prague culture are those of central and southeastern Transylvania, and they are linked to the social and cultural phenomena that are reflected in assemblages on the other side of the mountains, in Moldavia".

Curta (2018) continues that the sudden presence of handmade pottery and new settlement sites is a "good indication of migration", and that there's no consensus among archaeologists whether Slavs reached Moravia along Northern or Southern route, mountain slopes or rivers. Curta considered that it was a "forced movement of population done by and under the control of the Avars shortly after 600, perhaps in the circumstances surrounding the wars that Emperor Maurice was aggressively waging at that time across the Danube frontier of the empire against both Slavs and Avars", that there exist "more similarities between the archaeological record of northwestern Romania and Slovakia, on one hand, and those of southeastern Poland, on the other hand, than there are between Poland and Ukraine", but 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". There exists a scholarship contradiction in the interpretation of the archaeological data, and it is clear how much it's difficult and problematic to associate together with general culture, its remains, chronology, tribal identity, and population genetics.

3.1 Slavic tribes related to Korchak culture:

There exist many medieval Slavic tribes who were related to the Korchak culture and it's not an easy task to choose any one of them as PH908 carriers, but considering the cultural location, migration, and distribution of PH908, it probably had to be tribes which were at the same time present among both East and West Slavs and as such influence their ethnogenesis. In addition, it migrated to the South - such a description corresponds to 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 which various groups lived in Western Ukraine, the Southern Czech Republic, and the Middle Danube area between Lake Balaton and the Mur River, but besides that, there is anything much to say.

Croats (X within \) and Dulebes tribes (Volhynians, Drevlians, Polans within /), Sedov 1995
According to Sedov (1995), the White Croatian tribes emerged from Antes 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) criticized Sedov's consideration, which almost exclusively related White Croats with Penkovka culture and the Antes, because the territory White Croats inhabited in the middle and upper Dniester and the upper Vistula was part of Prague-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-Zhitomir culture, and near Southeastern Penkovka culture, the Early Slavs emigrated North, West, and South. Later in the 8th and 9th century from them developed the Luka-Raikovetskya culture to which belonged Eastern Polans, Drevlians, UlichsVolhynians, and Croats.

Image from "Русские" (1997), Russian Institute of Anthropology and Ethnography 
Both Sedov (1995) and Majorov (2006) with many other scholars agreed they were a large tribe, even a large Proto-Slavic tribe or among the oldest Slavic tribes, which got scattered in various directions during the Slavic migration period. 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".

Image from "Історія українського народу" (2010) by Семенюк Святослав Михайлович
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) or seven tribes and chieftains of Hungarian-Magyars (7) or 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 Sedov (1995), Korchinsky (2000, 2006) and others, with noted several gords and toponyms with Croatian ethnonym. In yellow color is lined the boundary of the territory of Lusatia where live the Sorbs, sometimes called the Lusatian Serbs (more about them below). 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.

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 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 high-quality interactive map see Scribble Maps
It is considered that they 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 which preserved Croatian ethnic identity as a hypothetical representative group for comparison of Y-DNA haplogroup distribution will be considered Rusyns and population of Western Ukraine, 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 CTS1211 subclade, and around 20% have I2, of which less than 5% 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 do not have well-defined Y-DNA subclades, but according to Mielnik-Sikorska et al. (2013), population of Lviv Oblast is 44.82% R1a, of which M458 is 13.65%, and 28.57% is I, while according to Utevska et al. (2015), almost the same percentages of R1a and I2-P37.2 (~30%) are found in Zakarpattia Oblast, while in Chernivtsi Oblast haplogroup I2 (>30%) occurs even more frequently than R1a-M198*. In comparison contemporary Croats mainly belong to subclades Z280 > CTS1211 (c. 20-25%) and I2 > PH908 (c. 30-35%). The lack of determined subclades limits the possibility of assumptions and conclusions, but if it is expected that the Croatian tribes were more similar to the "ancestral" population then seemingly exist some correlations between contemporary populations.

Sorbs in the Northwestern dotted area, Sedov 1995
According to DAI neighboring tribe to the White Croats in the West were 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 Eastern Galicia (see White Serbia#Dispute). While some scholars considered them only as local Slavs, Sedov (1995) argued that the medieval Sorbs would be a mixture of 1) local Slavs of the Tornovo cultural group of Sukow-Dziedzice culture 2) and migrant Slavs who did not have elements of Prague-Korchak culture yet middle Danube Avaro-Slav type of culture and that this migrant group has brought the S(o)erbian language and identity, and that contemporary Upper Sorbian dialect, which is spoken by the majority of Lusatian Sorbs, is related to that culture. According to Joachim Herrmann's Slavs, Avars and the Merovingian kingdom (1996), the 7th-century Frankish source by Fredegar mentions them in 630-631 as a tribe that was for a long time ruled by the Frankish kingdom, indicating that they settled the Saale-Elbe valley since the 6th century and the same is stated in DAI, and that unsuccessfully revolted against them during the Samo's union. According to DAI's origo gentis one brother took "half of the folk" and "came as the refugee to Heraclius".

Creative Commons
Since the ancestral ethnic group survived and preserved 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. 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 subclade M458, and only 4% have I2, of which 0% is PH908. In comparison, contemporary Serbs with R1a (c. 15-20%) mainly belong to subclade Z280 > CTS1211, as well have a high frequency of PH908 (c. 30-40%). Considering the historical facts it is expected that the Serbian tribes would have been more similar to the ancestral population, but that is not the case with the contemporary Serbs, and even if they did partly bring I2 > PH908 with them then that haplogroup would have been more present in contemporary Sorbs. These results imply that although share ethnic identity, happened ethnogenetic discontinuity between these two populations and that distribution of PH908, and a part of R1a, among the contemporary Serbs, mainly originates from a different route and population.

3.2 Slavic migrations to Southeastern Europe:

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 arrival did not finish by 8th 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).

Although there is no safe scientific evidence, in the late 5th and early 6th century possibly did happen first Slavic intrusions. 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 Antes and Pannonian Avars to the province Scythia Minor, and soon all over provinces of Thrace, Illyria, Dacia, Moesia, Dardania, Macedonia, Dalmatia, Epirus and so on.

Curta (2018) said 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", but, the archaeological and historical data and their interpretation are a bit contradicting.

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) is a representative of a migrationist model. According to archaeological data, he argued that in the Middle Danube basin was a mixture of Avaro-Slav Danube type of culture and Antes Penkovka type of culture, but also some Prague-Korchak culture, and from there migrated the main wave to the South; that exist rare findings of objects and ceramics of Prague-Korchak culture on the territory of Croatia, Bosnia and Herzegovina, and Serbia which were 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 a continuity with Slavic archaeological findings from the 8th-9th century. In conclusion, Sedov's arguing 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 it or were assimilated.

Dotted are Sukow-Dziedzice and Prague-Korchak cultures, Sedov 1995

On the other hand, Florin 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 ... but nothing indicates that the migrants were Slavs or that they spoke Slavic".

Croatian and Serbian tribes in the North and South, Sedov 1995
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".

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".

According to D. Dzino, The rise and fall of the Dalmatian ‘Big-men’: Social structures in Late Antique, Post-Roman and Early Medieval Dalmatia (ca. 500-850) (2014), the archaeological and historical interpretation sees a change in material culture through two frameworks – one implying settlement of the Croats as either small elite group or larger conglomeration of people and the other as the outcome of the contacts between local Slavs-Croats with the expanding Carolingian kingdom. The idea is that "we cannot understand post-Roman Dalmatia and the changes in 7th century without looking into changes in elite expression starting in the 5th and 6th century. In the same way, we cannot understand the formation of early medieval Dalmatia without looking into changes occurring in old-Croat cemeteries in the late 8th century". The "unification of burial rites coincided with the rise of dux Trepimirus (Trpimir, r. 840-850/864) and the elite-group who called themselves Croats as leaders of the Dalmatian duchy, regardless whether they were recent migrants, the existing local group, clan or newly forged elite-identity".

In the case of Serbs and Croats possibly happened a similar scenario as of the Hungarian and Bulgarian elite class. The poststructuralist perspective, although tends to overestimate the social-cultural influence, particularly explains such a process. 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 Danijel Dzino in Becoming Slav, Becoming Croat. Identity Transformations in Post-Roman and Early Medieval Dalmatia (2010) argue that ethnicities are not an immutable and static social phenomenon with the same core yet they are fluid. Population genetics and anthropology, without considering a multi-disciplinary approach even to an individual sample which results are often mistakenly extrapolated as representative of a collective group, give an answer to a biological category, while identity is a social category, and they should not be considered as equivalent through time in an attempt to prove a reflection of a contemporary nationalistic perspective to a distant period of time.

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 Eastern-Central route, lower percentage does not imply that the number of the Slavs was there less than among the Western South Slavs, on the contrary, in the hinterland of the territory of Western South Slavs most probably was a state of desolation which produced a founder effect of a small Slavic population resulting in an extreme lack of variance in both R1a and I2 haplogroups.

4. South Slavic languages:

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 doubts 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 South Slavic and East Slavic language family. At the time of arrival of Slavs in the 6th and early 7th century was spoken a variation of Proto-Slavic language and soon until the 9th century began a diversification. However, there still exist some doubts about the unity of South Slavic, but 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.

The South Slavic language group in its later diversification additionally branched into Western and Eastern. It is considered that within Western group never existed so-called "Middle South Slavic" (also known as Serbo-Croatian) which would have included Chakavian, Kajkavian and Shtokavian dialects, minus Slovenian language, because common innovations do not exist, rather, as other linguists have argued, from it branched five independent subbranches: Slovenian, Kajkavian, Chakavian, Western Shtokavian and Eastern Shokavian. Chakavian seemingly was never completely unified, Shtokavian from the early times was disunited with Western Shtokavian gravitating to Kajkavian while Eastern Shtokavian to Torlakian dialect.

Matasović (2008)
Mijo Lončarić in The Borders of the Kajkavian dialect in the past (1995) wrote about a common theory 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 the direction". Lončarić concluded that North of the Lake Balaton expanded the South Slavic language and that it did not happen any interruption there would have developed a new type of Kajkavian dialect. One can only wonder if this linguistic population is related to the Dulebes who lived between Lake Balaton and the Mur River.

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 favours a hypothesis that suggests an origin of the Slovenian language in the West Slavic language group (Bezlaj, 1967; Grafenauer, 1950; Žužek, 2007)". In short, the Slovenians who are genetically very close to West Slavs have a significantly lower frequency of M423 (~13%) and especially PH908 which separates them from other South Slavs and as such the Western route of migration for PH908 is historically and linguistically highly improbable.

On the general basis of the genetic relationship between languages, nevertheless, the exact date of their diversification, according to which the South Slavic and East Slavic are related, that Slovenian and Serbo-Croatian languages are related to Bulgarian and Macedonian languages and that Serbo-Croatian dialects are not related to Sorbian-Lusatian languages which on the contrary belong to West Slavic group - can be concluded that the majority of the Slavs arrived in Southeastern Europe from the Eastern rather the Western route, the Serbian and Croatian medieval tribes should have spoken the West Slavic language but as contemporary Serbo-Croatian dialects are not West Slavic languages these tribes did not leave enough linguistic trace and were assimilated by the majority, the contemporary Serbo-Croatian dialects which formed many centuries after the medieval migration cannot be used anyhow for the argumentation of PH908 migration with these tribes from the Western route which implies the PH908 mainly migrated with the other Slavs from the Eastern route, and the relationship of the languages support the ethnogenetic discontinuity between contemporary 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 region, approximately 30-60% of the native population. The Y-DNA haplogroup I2 > PH908 due to its young age and distribution is particularly taken into consideration. The presumed location of origination is around Western Ukraine where Early Slavs possibly assimilated the previous population. In the analysis arises potential cultural-tribal pattern Carpathian Tumuli culture > Chernyakhov culture > Korchak culture > Sclaveni > with White Croats and Dulebes as possible representatives of additional expansion of the haplogroup towards the West.

The autosomal DNA (Ralph et al. 2013), Y-DNA haplogroup R1a and I2 subclades prevalence, and archaeological data (Váňa 1983, Sedov 1995), mostly give undetermined resolutions but indicate an Eastern rather than the Western route of the main wave of Slavic migration, specifically in regard to carriers of PH908 subclades. Such a conclusion is also supported by a comparison of Y-DNA haplogroup frequency and variance in contemporary Serbs and Sorbs population, contemporary Croats 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) 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". To those Slavic tribes most probably also belonged so-called Seven Slavic tribes, including Severians, and others who became part of the First Bulgarian Empire. Besides that, the issue with the arrival of both Serbian and Croatian tribes is that according to DAI it was West of the Carpathians where is a lack of frequency and variance of PH908 in contemporary populations. While the Serbian tribes could only come from the Western route, the Croatian tribes could also come by Central and Eastern route both separately or as part of the main movement of the Slavs, however, there exists a historiographical dispute about the migration of Croatian tribes depending on the interpretation of DAI and other data.

However, archaeological data of the South Slavic migration, or even their lack of, has an on-going debate and various interpretations. Extreme lack of R1a and I2 subclades variance, especially in the Western Balkans even if is taken into account as precaution the post-migrational founder effect, does not significantly contradict poststructuralist perspective that there was no mass-migration yet a homogenous and seemingly small but large enough population to settle on a vastly depopulated territories where encountered a small and localized native population. Due to various ethnogenesis processes, including the fact the new Slavic identity earned prestige and its elite managed to claim a political rule over independent provinces, later kingdoms, the newcomer male population had an advantage over native males, which resulted with contemporary autosomal and Y-DNA distribution. The consequence of such a relationship, initially within the continent, is that the Late Middle Age Vlach tribe's male population in the Western Balkans often had Slavic PH908 haplogroup. In addition, among the South Slavic populations the matrix-nationalities, as well as languages, received the identity of Turkic-speaking Bulgars, and of Slavic-speaking Croats and Serbs, in both cases most probably only minority groups, or in the latter case even as significantly numerous groups, that were imposed as a ruling class with state-building skills.