Slavic speaking populations are the most numerous Indo-European ethno linguistic group in Europe. They show great variety and fall into three groups: West, East and South Slavic populations. In order to contribute to the understanding of the correlation between linguistic and genetic affiliation of Slavic populations, we have analyzed for the first time their matrilineal and patrilineal relationships and we have also illustrated their position in the European uniparental genetic landscape. For the purpose, we have collected previously published data for the frequencies of mitochondrial DNA (mtDNA) and Y-chromosome haplogroups in Slavic and other European populations and compared them by Principal Component Analysis (PCA). In the inter-Slavic population comparisons, West and East Slavs are in a closer position, whereas South Slavic populations are rather grouped on their own. In the European context, South Slavic populations are positioned more close to neighboring Balkan non-Slavic and North Italian populations, than to other Slavic populations. When considering the uniparental diversity of Slavic speaking populations, one should also take into account the prevalence of Y-chromosome haplogroup N among East Slavs (comprising almost half of the paternal gene pool in instances), which is almost absent among the other groups (not exceeding 2% - 3%). In conclusion, the data in the present study point that West-East and South Slavic speaking populations, behave as separate groups based on their uniparental genetic structure, which shows that they do not share substantial common genetic ancestry and that there is great genetic variety in the Slavic linguistic unity.
Slavs are the most numerous Indo-European ethno-linguistic group in Europe ( Kipfer, 2000 ). Their proposed homeland is in the middle Dnieper basin: the area North-East of the Carpathians, the upper stream of rivers Bug and Dniester, mainly around Pripet river ( Mutafchiev, 1943 ; Rebala et al., 2007 ). They spread from the 6th century AD to inhabit whole Eastern Europe and parts of Central and South-Eastern Europe.
Roman writers mentioned Slavs as Venethi or Venedi ( Curta, 2001 ). The Southern European Slavs are first named as Sclavenes by East Roman (Byzantine) authors in the 6th century AD ( Džino, 2010 ; Smith, 2005 ).
Extant Slavic languages show great variety and fall into three groups: West (Czech, Slovakian and Polish), East (Russian, Belorussian and Ukrainian) and South (Serbian, Croatian, Bulgarian and Slovenian). The Bulgarian literary language differs from other Slavic languages by the almost complete loss of grammatical case; the creation of definite article of nouns (appearing in the form of a suffix, added to the stem); analytical comparative and superlative (by word-particles); and a complex tense system where the infinitive is completely lost ( Aepli, von Waldenfels, & Samardzic, 2014 ; Kushniarevich et al., 2015 ; Raykov, 2005 ).
Orthodox Christian Slavs use the Cyrillic alphabet, while Roman Catholic Slavs and Bosniaks use the Latin alphabet.
Genetic and genomic analysis of Slavs from different countries have been the object of many previously published studies; however, comparative genetic investigations among different Slavic groups are only few ( Grzybowski et al., 2007 ; Malyarchuk et al., 2008 ; Mielnik-Sikorska et al., 2013 ; Rebala et al., 2007 ). To the best of our knowledge, the most comprehensive study of Slavic genetic heritage to date ( Kushniarevich et al., 2015 ) reveals that the Slavic genetic diversity was formed through assimilation of preexisting regional genetic components and in situ gene pool shaping; as it also identifies an apparent genetic homogeneity of the majority of West and East Slavs and a substantial genetic difference between them and South Slavs.
In order to further contribute to the understanding of the correlation between language and genetic origin in Slavs, the present study analyzes for the first time the matrilineal and patrilineal relationships among European Slavic-speaking countries’ populations and also illustrates their position in the European uniparental genetic landscape.
We have collected previously published data for the frequencies of mitochondrial DNA (mtDNA) and Y-chromosome haplogroups in Slavic speaking and other European populations. In these studies, the mtDNA haplogroup assignment was based on partial or entire control region sequences and/or coding region markers; and the Y-chromosome haplogroup classification was performed by genotyping of informative biallelic markers. To analyze comparable results for larger number of populations, the data were normalized to the highest possible level of phylogenetic resolution.
We have included 18 and 27 (sub-) populations in the mtDNA and Y-chro- mosome haplogroup analyses of Slavic speaking populations (Supplementary
The mtDNA and Y-chromosome relationships among the populations were depicted by Principal Component Analysis (PCA) performed using XLSTAT.
The plot of the PCA performed on mtDNA haplogroup frequencies in Slavic speaking populations is presented in
Patrilineal relationships of Slavic speaking populations based on Y-chromo- some haplogroup frequencies are depicted in the PCA plot in
In the PC analyses of both mtDNA and Y-chromosome haplogroup frequencies of Slavic speaking populations, South Slavs are quite dispersed. This is probably due to their different pasts. It is established that the ancestors of Croatians and Serbians migrated to the Balkan Peninsula in the 8th century AD from territories in Central Europe (White Croats and White Serbs) ( Borri, 2011 ; Chadwick, 2014 ). The area of their dispersal is in the territory of present-day West Slavic populations’ countries. Very different is the story of the contemporary Bulgarians. South Slavic tribes (Sclavеnes) and Proto-Bulgarians arrived almost simultaneously on the Balkan Peninsula in the 7th century AD, when the
Proto-Bulgarians established the Danubian Bulgarian state ( Dobrev, 2016 ; Mutafchiev, 1943 ).
The position of the mtDNA haplogroup frequency profiles of Slavic speaking populations in the European context is represented in
The comparison of the Y-chromosome haplogroup frequencies in Slavic speaking and remaining European populations performed by PC analysis is given in
In general, the obtained results show that based on the distribution of mtDNA and Y-chromosome haplogroups West and East Slavic speaking populations locate separately from South Slavic populations. Furthermore, in the European uniparental landscape South Slavic speaking populations are positioned more close to neighboring Balkan non-Slavic populations and North Italian populations, than to other Slavic populations. This hints that the linguistic resemblance of South Slavic speaking populations with East and West Slavic groups is not paralleled to a similar extent by a genetic one, which is in line with previous findings demonstrating that the basis of the gene pool of West-East and South Slavic speaking populations is different ( Kushniarevich et al., 2015 ).
When considering the uniparental diversity of Slavic speaking populations, one should highlight a peculiarity in the East Slavs’ gene pool, namely the prevalence of Y-chromosome haplogroup N, which is typical for certain Asian populations: China-Naxi―25%, China-Oroqen―28.5%, China-Tu―28.5%, Cam- bodians―16.7% ( Sengupta et al., 2006 ), Mongols―8.7% ( Derenko et al., 2007 ; Hammer et al., 2005 ), Manchurian―4.9% ( Hammer et al., 2005 ), Koreans―4.3% ( Hammer et al., 2005 ; Zhong et al., 2010 ), Vietnamese―2.8% ( Hammer et al., 2005 ); Central Asian Turkic populations, such as Khakassians―50% and
Altaians―9.2% ( Derenko et al., 2007 ); and several Siberian populations: Khanty― 81.5% ( Mirabal et al., 2009 ), Yakuts―80.0% ( Derenko et al., 2007 ), Buryats― 30.9% ( Hammer et al., 2005 ), Evenks―29.3%, Tuvininas―29.1% and Koryaks― 25% ( Derenko et al., 2007 ).
In Europe Y-chromosome haplogroup N shows high frequency in Saami― 44.7% ( Karlsson et al., 2006 ). The frequency values for European Russians are striking: Arkhangel―41.1% ( Balanovsky et al., 2008 ; Mirabal et al., 2009 ), Vologda―38.8%, Pskov―24.2%, Smolensk―14.0%, Kostroma―13.5% ( Balano- vsky et al., 2008 ), Kursk―12.9% ( Balanovsky et al., 2008 ; Mirabal et al., 2009 ), Belgorod―12.6% ( Balanovsky et al., 2008 ), Tver―11.7% ( Balanovsky et al., 2008 ; Mirabal et al., 2009 ), Voronezh―6.3%, Orel province―5.5% ( Balanovsky et al., 2008 ) and Kalmyks―3.3% ( Derenko et al., 2007 ). The data for Bela- russians are 9.6% ( Kushniarevich et al., 2013 ; Kushniarevich et al., 2015 ) and for Ukrainians―5.7% ( Battaglia et al., 2009 ; Kushniarevich et al., 2015 ).
In the Y-chromosome gene pool of any West or South Slavic speaking population haplogroup N does not exceed 2-3% in frequency: Poles―2.7% ( Battaglia et al., 2009 ; Rebala et al., 2013 ), Czechs―2.7%, Croats―1.7% ( Battaglia et al., 2009 ), Bulgarians―0.5% ( Karachanak et al., 2013 ), Serbs 1.9% ( Regueiro et al., 2012 ), Slovenians―0% and Bosniaks―0% ( Battaglia et al., 2009 ). The same pattern is observed in the remaining populations in Europe, the Caucasus: Balkarians―0% and Georgians―0% ( Battaglia et al., 2009 ) and in South Asia ( Sengupta, et al., 2006 ). Obviously, this characteristic in the patrilineal gene pool of East Slavs can be explained by the Mongol invasion and the presence of Mongols in Russia (called Tatars by Russians) between 1237 and 1480 AD ( Karamzin, 1811/2016 ; Deynichenko, 2003 ).
In conclusion, as illustrated by the PC analysis of mtDNA and Y-chromosome haplogroup frequencies, West-East and South Slavic speaking populations, traditionally called “Slavs” (a term introduced in the 16th century AD) ( Šafařik, 1848 ) are heterogeneous based on the uniparental genetic diversity, which shows that they do not share substantial common genetic ancestry and that there is great genetic variety in the Slavic linguistic unity.
We would like to thank prof. Antonio Torroni, prof. Ornella Semino and their collaborators (Department of Biology and Biotechnology, University of Pavia, Italy) for their contribution for the development of the uniparental genetic study in Bulgaria.
Karachanak-Yan- kova, S., Nesheva, D., Toncheva, D., & Galabov, A. S. (2017). The Uniparental Genetic Landscape of Modern Slavic Speaking Populations. Advances in Anthropology, 7, 318-332. https://doi.org/10.4236/aa.2017.74018
Population | Total | H | HV | J | T | U1 | U2 | U3 | U4 | U5a | U5b | U6 | U7 | U8 | U* | K | N1 | N2 | X | M | L |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Belarusa | 267 | 99 | 25 | 24 | 20 | 1 | 4 | 6 | 10 | 17 | 15 | 0 | 1 | 0 | 0 | 10 | 15 | 9 | 3 | 7 | 0 |
Bosniab,c | 383 | 171 | 37 | 28 | 15 | 3 | 2 | 3 | 18 | 22 | 14 | 1 | 4 | 1 | 4 | 22 | 12 | 11 | 7 | 2 | 1 |
Bulgariansd | 885 | 359 | 65 | 67 | 91 | 12 | 12 | 16 | 33 | 37 | 20 | 0 | 4 | 4 | 0 | 50 | 22 | 24 | 18 | 8 | 3 |
Croatiac | 569 | 255 | 43 | 58 | 41 | 6 | 19 | 9 | 13 | 33 | 18 | 1 | 2 | 1 | 5 | 26 | 17 | 10 | 9 | 0 | 1 |
Czechse | 179 | 79 | 7 | 21 | 22 | 0 | 1 | 4 | 5 | 9 | 6 | 0 | 0 | 1 | 0 | 7 | 8 | 1 | 3 | 3 | 0 |
Herzegovinac | 130 | 55 | 14 | 11 | 8 | 2 | 3 | 0 | 6 | 3 | 3 | 0 | 0 | 0 | 0 | 12 | 3 | 3 | 1 | 2 | 0 |
Kalugaf | 71 | 27 | 4 | 3 | 10 | 1 | 2 | 1 | 6 | 6 | 2 | 0 | 0 | 0 | 0 | 3 | 1 | 0 | 2 | 2 | 1 |
Polandg | 413 | 174 | 25 | 32 | 31 | 2 | 7 | 4 | 21 | 31 | 20 | 0 | 0 | 5 | 0 | 18 | 15 | 15 | 8 | 2 | 0 |
Pskovf | 68 | 29 | 4 | 8 | 4 | 0 | 2 | 0 | 2 | 12 | 2 | 0 | 0 | 1 | 0 | 0 | 0 | 2 | 0 | 1 | 0 |
Serbiansh | 139 | 66 | 9 | 9 | 7 | 1 | 1 | 2 | 12 | 5 | 4 | 0 | 1 | 0 | 0 | 11 | 3 | 3 | 2 | 1 | 1 |
Slovakse | 207 | 94 | 14 | 19 | 22 | 1 | 2 | 0 | 11 | 4 | 12 | 0 | 1 | 0 | 0 | 3 | 11 | 6 | 3 | 2 | 2 |
Sloveniab,c | 201 | 86 | 10 | 24 | 21 | 1 | 5 | 3 | 7 | 14 | 10 | 0 | 0 | 1 | 0 | 9 | 2 | 6 | 2 | 0 | 0 |
Tulaf | 73 | 43 | 3 | 4 | 7 | 0 | 0 | 2 | 3 | 2 | 2 | 0 | 0 | 0 | 0 | 2 | 4 | 0 | 0 | 0 | 1 |
Ukrainiansi | 607 | 233 | 49 | 50 | 72 | 0 | 9 | 6 | 35 | 36 | 24 | 0 | 0 | 1 | 0 | 30 | 25 | 16 | 8 | 7 | 1 |
Velikii Novgorodg | 79 | 33 | 6 | 4 | 5 | 0 | 2 | 1 | 0 | 12 | 3 | 0 | 0 | 3 | 0 | 0 | 3 | 1 | 1 | 2 | 0 |
Vladimirf | 72 | 25 | 1 | 6 | 14 | 0 | 1 | 1 | 2 | 5 | 0 | 0 | 1 | 0 | 0 | 5 | 6 | 3 | 1 | 1 | 0 |
Volotg | 78 | 32 | 10 | 7 | 7 | 0 | 0 | 0 | 1 | 3 | 1 | 0 | 0 | 2 | 0 | 3 | 2 | 2 | 0 | 5 | 0 |
Yaroslavlf | 41 | 20 | 3 | 2 | 1 | 0 | 0 | 0 | 2 | 4 | 0 | 0 | 0 | 0 | 0 | 3 | 3 | 0 | 2 | 1 | 0 |
a Kushniarevich et al., 2013 ; bMalyarchuk et al., 2003; cSarac et al., 2014;dKarachanak et al., 2012; e Malyarchuk et al., 2008 ; fMalyarchuk et al., 2004; g Grzybowsky et al., 2007 ; hDavidovic et al., 2015; iPshenichnov et al., 2013.
Population | Total | C-M217 | E-M78 | G-P15 | G-M285 | I-M253 | I-P37 | I-M223 | I-M170 (x M253, P37, M223) | J-M304 (x M172) | J-M172 | N-M231 | T-M70 | Q-M242 | R-SRY1532 | R-M343 (x M269) | R-M269 | R-M124 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Arkhangela,b | 287 | 0 | 1 | 2 | 0 | 16 | 14 | 0 | 13 | 1 | 5 | 118 | 0 | 1 | 92 | 0 | 22 | 0 |
Belarus Centrec | 88 | 0 | 1 | 1 | 0 | 6 | 16 | 3 | 0 | 1 | 0 | 7 | 0 | 0 | 44 | 0 | 7 | 1 |
Belarus Eastc,d | 182 | 0 | 7 | 1 | 0 | 2 | 29 | 1 | 1 | 2 | 3 | 13 | 1 | 0 | 106 | 0 | 14 | 0 |
Belarus East Polesiec | 96 | 0 | 1 | 6 | 0 | 4 | 25 | 0 | 0 | 2 | 4 | 8 | 1 | 0 | 41 | 0 | 2 | 0 |
Belarus Northc,d | 201 | 0 | 6 | 0 | 0 | 3 | 26 | 4 | 0 | 1 | 5 | 25 | 0 | 0 | 119 | 0 | 12 | 0 |
Belarus Westc | 73 | 0 | 4 | 0 | 0 | 5 | 2 | 1 | 0 | 1 | 1 | 10 | 0 | 0 | 42 | 1 | 6 | 0 |
Belarus West Polesiec | 121 | 0 | 2 | 1 | 0 | 10 | 31 | 0 | 0 | 1 | 4 | 10 | 1 | 1 | 54 | 1 | 5 | 0 |
Belgoroda | 143 | 1 | 1 | 1 | 0 | 5 | 18 | 1 | 0 | 0 | 6 | 18 | 2 | 0 | 85 | 0 | 4 | 0 |
Bosnia-Croatse | 90 | 0 | 8 | 1 | 0 | 0 | 66 | 0 | 0 | 0 | 2 | 0 | 0 | 0 | 11 | 0 | 2 | 0 |
Bosnianse | 84 | 0 | 11 | 3 | 0 | 4 | 38 | 0 | 0 | 2 | 10 | 0 | 0 | 0 | 13 | 0 | 3 | 0 |
Bosnia-Serbse | 81 | 0 | 16 | 1 | 0 | 2 | 28 | 1 | 2 | 1 | 7 | 5 | 0 | 0 | 11 | 0 | 5 | 0 |
Bulgariansf | 808 | 4 | 158 | 40 | 0 | 35 | 163 | 14 | 3 | 27 | 85 | 4 | 13 | 4 | 138 | 2 | 86 | 1 |
Croatse | 118 | 0 | 11 | 5 | 0 | 8 | 38 | 1 | 0 | 0 | 7 | 2 | 0 | 0 | 35 | 0 | 11 | 0 |
Czechse | 75 | 0 | 0 | 3 | 0 | 7 | 0 | 1 | 0 | 0 | 4 | 2 | 1 | 0 | 31 | 1 | 25 | 0 |
Kostromaa | 52 | 0 | 3 | 0 | 0 | 6 | 6 | 2 | 0 | 2 | 2 | 7 | 0 | 0 | 17 | 1 | 5 | 0 |
Kurska,b | 85 | 0 | 5 | 0 | 0 | 4 | 11 | 1 | 0 | 0 | 2 | 11 | 1 | 0 | 46 | 0 | 4 | 0 |
Orela | 110 | 0 | 1 | 0 | 0 | 9 | 15 | 1 | 0 | 0 | 1 | 6 | 3 | 1 | 69 | 0 | 4 | 0 |
Polesa,g | 619 | 0 | 17 | 7 | 1 | 56 | 28 | 9 | 14 | 2 | 12 | 17 | 0 | 3 | 371 | 0 | 80 | 0 |
Pskova | 132 | 0 | 4 | 0 | 0 | 7 | 13 | 0 | 2 | 1 | 1 | 32 | 0 | 0 | 64 | 0 | 5 | 0 |
Serbsh | 103 | 0 | 18 | 6 | 0 | 8 | 30 | 1 | 0 | 1 | 7 | 2 | 0 | 0 | 21 | 0 | 8 | 0 |
Slovakia Westerng | 164 | 0 | 6 | 8 | 0 | 9 | 25 | 3 | 0 | 3 | 3 | 5 | 1 | 0 | 74 | 0 | 25 | 0 |
Slovenianse | 75 | 0 | 2 | 2 | 0 | 5 | 15 | 2 | 1 | 1 | 2 | 0 | 0 | 0 | 29 | 0 | 16 | 0 |
Smolenska | 107 | 1 | 8 | 0 | 0 | 2 | 11 | 0 | 0 | 0 | 3 | 15 | 3 | 0 | 49 | 0 | 12 | 0 |
Tvera,b | 111 | 0 | 3 | 0 | 0 | 5 | 10 | 3 | 0 | 0 | 5 | 13 | 0 | 1 | 63 | 0 | 6 | 0 |
Ukrainiansd,e | 748 | 1 | 49 | 21 | 0 | 38 | 145 | 2 | 17 | 3 | 24 | 43 | 9 | 3 | 338 | 1 | 49 | 0 |
Vologdaa | 121 | 0 | 1 | 1 | 2 | 14 | 10 | 2 | 0 | 0 | 1 | 47 | 0 | 2 | 40 | 0 | 1 | 0 |
Voronezha | 96 | 0 | 1 | 2 | 0 | 3 | 16 | 0 | 1 | 2 | 1 | 6 | 0 | 0 | 57 | 0 | 5 | 1 |
a Balanovsky et al., 2008 ; b Mirabal et al., 2009 ; c Kushniarevich et al., 2013 ; d Kushniarevich et al., 2015 ; e Battaglia et al., 2009 ; f Battaglia et al., 2009 ; g Rebala et al., 2013 ; hReguiero et al., 2012.
Population | Total | H | HV | J | T | U1 | U2 | U3 | U4 | U5a | U5b | U6 | U7 | U8 | U* | K | N1 | N2 | X | M | L |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Andalusiansa | 279 | 113 | 17 | 19 | 21 | 0 | 2 | 8 | 3 | 4 | 21 | 16 | 1 | 0 | 1 | 20 | 4 | 8 | 6 | 1 | 10 |
Austriab | 273 | 119 | 7 | 23 | 36 | 2 | 6 | 3 | 13 | 16 | 9 | 0 | 0 | 0 | 3 | 23 | 4 | 4 | 4 | 1 | 0 |
Belarus Centralc | 47 | 20 | 6 | 4 | 1 | 0 | 0 | 1 | 2 | 6 | 1 | 0 | 1 | 0 | 0 | 0 | 3 | 1 | 1 | 0 | 0 |
Belarus Eastc | 87 | 31 | 9 | 5 | 8 | 0 | 3 | 2 | 2 | 5 | 6 | 0 | 0 | 0 | 0 | 4 | 6 | 3 | 1 | 2 | 0 |
Belarus Northc | 55 | 20 | 1 | 9 | 5 | 1 | 0 | 0 | 3 | 4 | 2 | 0 | 0 | 0 | 0 | 2 | 1 | 3 | 1 | 2 | 0 |
Belarus Westc | 78 | 28 | 9 | 6 | 6 | 0 | 1 | 3 | 3 | 2 | 6 | 0 | 0 | 0 | 0 | 4 | 5 | 2 | 0 | 3 | 0 |
Bolognad | 35 | 16 | 1 | 5 | 3 | 0 | 0 | 1 | 0 | 2 | 0 | 0 | 0 | 1 | 0 | 3 | 1 | 0 | 1 | 0 | 0 |
Bosniae,f | 383 | 171 | 37 | 28 | 15 | 3 | 2 | 3 | 18 | 22 | 14 | 1 | 4 | 1 | 4 | 22 | 12 | 11 | 7 | 2 | 1 |
Bulgariansg | 885 | 359 | 65 | 67 | 91 | 12 | 12 | 16 | 33 | 37 | 20 | 0 | 4 | 4 | 0 | 50 | 22 | 24 | 18 | 8 | 3 |
Center Portugalh | 78 | 47 | 0 | 6 | 5 | 0 | 1 | 1 | 0 | 1 | 2 | 3 | 0 | 0 | 0 | 7 | 0 | 1 | 0 | 0 | 4 |
Croatiaf | 569 | 255 | 43 | 58 | 41 | 6 | 19 | 9 | 13 | 33 | 18 | 1 | 2 | 1 | 5 | 26 | 17 | 10 | 9 | 0 | 1 |
Czechsi | 179 | 79 | 7 | 21 | 22 | 0 | 1 | 4 | 5 | 9 | 6 | 0 | 0 | 1 | 0 | 7 | 8 | 1 | 3 | 3 | 0 |
Finlandj | 200 | 80 | 9 | 9 | 12 | 0 | 1 | 0 | 4 | 13 | 36 | 0 | 1 | 0 | 0 | 13 | 7 | 5 | 5 | 0 | 0 |
Francek | 173 | 74 | 5 | 11 | 22 | 2 | 5 | 0 | 2 | 9 | 4 | 2 | 0 | 0 | 3 | 22 | 5 | 3 | 3 | 0 | 0 |
Galicia (Northwestern Spain)h | 43 | 22 | 4 | 1 | 3 | 0 | 2 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 4 | 0 | 1 | 1 | 0 | 4 |
Germanyl | 213 | 90 | 0 | 20 | 26 | 1 | 1 | 1 | 12 | 10 | 6 | 0 | 0 | 0 | 3 | 14 | 4 | 7 | 2 | 1 | 0 |
Herzegovinaf | 130 | 55 | 14 | 11 | 8 | 2 | 3 | 0 | 6 | 3 | 3 | 0 | 0 | 0 | 0 | 12 | 3 | 3 | 1 | 2 | 0 |
Kalugam | 71 | 27 | 4 | 3 | 10 | 1 | 2 | 1 | 6 | 6 | 2 | 0 | 0 | 0 | 0 | 3 | 1 | 0 | 2 | 2 | 1 |
La Spezia and Tuscanyd | 52 | 29 | 2 | 3 | 4 | 0 | 0 | 0 | 1 | 1 | 0 | 2 | 0 | 0 | 0 | 7 | 2 | 0 | 0 | 0 | 0 |
North Portugalh | 84 | 39 | 4 | 2 | 15 | 0 | 1 | 2 | 2 | 1 | 3 | 1 | 0 | 0 | 0 | 4 | 5 | 4 | 0 | 0 | 1 |
Northeast Italyd | 108 | 42 | 8 | 6 | 16 | 1 | 1 | 1 | 2 | 6 | 5 | 0 | 0 | 0 | 0 | 9 | 5 | 4 | 0 | 2 | 0 |
Northern Greeksn | 319 | 133 | 13 | 33 | 27 | 6 | 1 | 12 | 14 | 12 | 3 | 2 | 2 | 0 | 0 | 16 | 11 | 8 | 12 | 4 | 0 |
Northwest Italyd | 162 | 73 | 14 | 19 | 12 | 1 | 5 | 4 | 3 | 7 | 1 | 0 | 0 | 0 | 0 | 10 | 5 | 2 | 3 | 1 | 0 |
Poland North˚ | 253 | 109 | 18 | 20 | 25 | 0 | 3 | 1 | 15 | 14 | 12 | 0 | 0 | 3 | 0 | 8 | 11 | 8 | 4 | 1 | 0 |
Poland Northeast˚ | 73 | 27 | 4 | 6 | 3 | 0 | 0 | 1 | 1 | 12 | 3 | 0 | 0 | 0 | 0 | 7 | 3 | 2 | 2 | 0 | 0 |
Poland South˚ | 87 | 38 | 3 | 6 | 3 | 2 | 4 | 2 | 5 | 5 | 5 | 0 | 0 | 2 | 0 | 3 | 1 | 5 | 2 | 1 | 0 |
Pskovm | 68 | 29 | 4 | 8 | 4 | 0 | 2 | 0 | 2 | 12 | 2 | 0 | 0 | 1 | 0 | 0 | 0 | 2 | 0 | 1 | 0 |
Romaniap | 128 | 67 | 9 | 8 | 8 | 0 | 1 | 0 | 7 | 6 | 2 | 0 | 0 | 1 | 0 | 7 | 0 | 7 | 1 | 1 | 0 |
Sardiniad | 70 | 32 | 3 | 9 | 11 | 0 | 0 | 0 | 0 | 2 | 5 | 0 | 0 | 1 | 0 | 5 | 2 | 0 | 0 | 0 | 0 |
Serbiansq | 139 | 66 | 9 | 9 | 7 | 1 | 1 | 2 | 12 | 5 | 4 | 0 | 1 | 0 | 0 | 11 | 3 | 3 | 2 | 1 | 1 |
Slovaksh | 207 | 94 | 14 | 19 | 22 | 1 | 2 | 0 | 11 | 4 | 12 | 0 | 1 | 0 | 0 | 3 | 11 | 6 | 3 | 2 | 2 |
Sloveniae,f | 201 | 86 | 10 | 24 | 21 | 1 | 5 | 3 | 7 | 14 | 10 | 0 | 0 | 1 | 0 | 9 | 2 | 6 | 2 | 0 | 0 |
South Portugalh | 137 | 66 | 4 | 11 | 9 | 0 | 1 | 1 | 3 | 3 | 2 | 0 | 0 | 0 | 0 | 8 | 1 | 4 | 5 | 2 | 17 |
Sweden (Saami excluded)r | 284 | 134 | 13 | 27 | 23 | 1 | 4 | 0 | 8 | 14 | 11 | 0 | 2 | 2 | 0 | 18 | 8 | 4 | 7 | 4 | 4 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Swedish Saamir | 37 | 0 | 25 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 12 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Tulam | 73 | 43 | 3 | 4 | 7 | 0 | 0 | 2 | 3 | 2 | 2 | 0 | 0 | 0 | 0 | 2 | 4 | 0 | 0 | 0 | 1 |
Ukrainianss | 607 | 233 | 49 | 50 | 72 | 0 | 9 | 6 | 35 | 36 | 24 | 0 | 0 | 1 | 0 | 30 | 25 | 16 | 8 | 7 | 1 |
VelikiiNovgorod˚ | 79 | 33 | 6 | 4 | 5 | 0 | 2 | 1 | 0 | 12 | 3 | 0 | 0 | 3 | 0 | 0 | 3 | 1 | 1 | 2 | 0 |
Vladimirm | 72 | 25 | 1 | 6 | 14 | 0 | 1 | 1 | 2 | 5 | 0 | 0 | 1 | 0 | 0 | 5 | 6 | 3 | 1 | 1 | 0 |
Volot˚ | 78 | 32 | 10 | 7 | 7 | 0 | 0 | 0 | 1 | 3 | 1 | 0 | 0 | 2 | 0 | 3 | 2 | 2 | 0 | 5 | 0 |
Yaroslavlm | 41 | 20 | 3 | 2 | 1 | 0 | 0 | 0 | 2 | 4 | 0 | 0 | 0 | 0 | 0 | 3 | 3 | 0 | 2 | 1 | 0 |
a Hernаndez et al., 2014; bBrandstätter et al., 2007; c Kushniarevich et al., 2013 ; dBoattini et al., 2013; eMalyarchuk et al., 2003; fSarac et al., 2014; gKarachanak et al., 2012; hGonzález et al., 2003; i Malyarchuk et al., 2008 ; jHedman et al., 2007; kRichard et al., 2007; lTetzlaff et al., 2007; mMalyarchuk et al., 2004; nIrwin et al., 2008; o Grzybowsky et al., 2007 ; pHervella et al., 2014; qDavidovic et al., 2015; rTillmar et al., 2010; sPshenichnov et al., 2013.
Population | Total | Е-М35 | G-M201 | I-M253 | I-P37 | I-M223 | I-M170 (x M253, P37, M223) | J-M304 | N-M231 | T-M70 | Q-M242 | R-SRY1532 | R-M343 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Arkhangela,b | 287 | 1 | 2 | 16 | 14 | 0 | 13 | 6 | 118 | 0 | 1 | 92 | 22 |
Belarusc,d | 761 | 24 | 9 | 30 | 129 | 9 | 1 | 25 | 73 | 3 | 1 | 406 | 48 |
Belgoroda | 143 | 1 | 1 | 5 | 18 | 1 | 0 | 6 | 18 | 2 | 0 | 85 | 4 |
Bolognae | 29 | 3 | 3 | 3 | 0 | 0 | 0 | 3 | 0 | 0 | 0 | 0 | 17 |
Bosnia-Croatsf | 90 | 8 | 1 | 0 | 66 | 0 | 0 | 2 | 0 | 0 | 0 | 11 | 2 |
Bosniansf | 84 | 11 | 3 | 4 | 38 | 0 | 0 | 12 | 0 | 0 | 0 | 13 | 3 |
Bosnia-Serbsf | 81 | 18 | 1 | 2 | 28 | 1 | 2 | 8 | 5 | 0 | 0 | 11 | 5 |
Bulgariansg | 808 | 175 | 40 | 35 | 163 | 14 | 3 | 112 | 4 | 13 | 4 | 138 | 88 |
Cataloniah | 2309 | 191 | 100 | 44 | 67 | 37 | 11 | 266 | 0 | 28 | 0 | 32 | 1525 |
Cretei | 193 | 17 | 21 | 8 | 3 | 3 | 11 | 75 | 0 | 4 | 0 | 17 | 33 |
Croatsf | 118 | 11 | 5 | 8 | 38 | 1 | 0 | 7 | 2 | 0 | 0 | 35 | 11 |
Czechsf | 75 | 0 | 3 | 7 | 0 | 1 | 0 | 4 | 2 | 1 | 0 | 31 | 26 |
Finland-Osterbottenj | 40 | 0 | 2 | 8 | 0 | 0 | 0 | 0 | 26 | 0 | 0 | 3 | 1 |
Germany-Bavariak | 218 | 18 | 7 | 37 | 6 | 6 | 0 | 16 | 2 | 2 | 1 | 27 | 95 |
Germany-Mecklenburgk | 131 | 7 | 2 | 19 | 3 | 7 | 1 | 4 | 3 | 1 | 0 | 18 | 66 |
Greeksf,i | 149 | 43 | 6 | 4 | 10 | 2 | 1 | 33 | 0 | 5 | 0 | 21 | 21 |
Hungarians f,l | 245 | 18 | 11 | 16 | 35 | 5 | 2 | 21 | 1 | 1 | 1 | 81 | 42 |
Italy Centrale | 77 | 8 | 12 | 1 | 0 | 1 | 1 | 21 | 0 | 0 | 0 | 3 | 29 |
Italy Northeaste | 73 | 11 | 5 | 8 | 0 | 0 | 1 | 11 | 0 | 2 | 0 | 0 | 28 |
Italy Northweste | 161 | 19 | 13 | 5 | 1 | 1 | 6 | 17 | 0 | 2 | 0 | 5 | 89 |
Italy Southe | 198 | 35 | 34 | 7 | 4 | 2 | 1 | 42 | 0 | 9 | 0 | 8 | 51 |
Kurska,b | 85 | 5 | 0 | 4 | 11 | 1 | 0 | 2 | 11 | 1 | 0 | 46 | 4 |
La Spezia and Tuscanye | 123 | 14 | 8 | 5 | 1 | 1 | 0 | 14 | 0 | 4 | 0 | 6 | 67 |
Livnia | 110 | 1 | 0 | 9 | 15 | 1 | 0 | 1 | 6 | 3 | 1 | 69 | 4 |
Macedonian Greeksf,i | 114 | 24 | 3 | 6 | 15 | 1 | 2 | 21 | 0 | 2 | 0 | 19 | 19 |
Moldaviansm | 125 | 16 | 1 | 6 | 26 | 4 | 0 | 10 | 2 | 1 | 1 | 38 | 20 |
Ostrova | 75 | 3 | 0 | 5 | 7 | 0 | 1 | 2 | 21 | 0 | 0 | 34 | 2 |
Polesf,k | 619 | 19 | 8 | 56 | 28 | 9 | 14 | 14 | 17 | 0 | 3 | 371 | 80 |
Porhova | 57 | 2 | 0 | 2 | 6 | 0 | 1 | 0 | 11 | 0 | 0 | 30 | 3 |
Repievkaa | 96 | 1 | 2 | 3 | 16 | 0 | 1 | 3 | 6 | 0 | 0 | 57 | 5 |
Romaniansl,m | 232 | 39 | 7 | 12 | 53 | 5 | 5 | 36 | 1 | 1 | 1 | 46 | 25 |
Roslavla | 107 | 8 | 0 | 2 | 11 | 0 | 0 | 3 | 15 | 3 | 0 | 49 | 12 |
Sardiniae | 82 | 8 | 11 | 0 | 32 | 2 | 0 | 8 | 0 | 1 | 0 | 0 | 18 |
Serbsn | 103 | 19 | 6 | 8 | 30 | 1 | 0 | 8 | 2 | 0 | 0 | 21 | 8 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Sicilye | 141 | 25 | 17 | 2 | 2 | 2 | 1 | 36 | 0 | 3 | 0 | 8 | 43 |
Slovakia Westernk | 164 | 7 | 8 | 9 | 25 | 3 | 0 | 6 | 5 | 1 | 0 | 74 | 25 |
Sloveniansf | 75 | 2 | 2 | 5 | 15 | 2 | 1 | 3 | 0 | 0 | 0 | 29 | 16 |
Swedish Saamij | 38 | 0 | 0 | 12 | 0 | 0 | 0 | 0 | 17 | 0 | 0 | 6 | 3 |
Tvera,b | 111 | 4 | 0 | 5 | 10 | 3 | 0 | 5 | 13 | 0 | 1 | 63 | 6 |
Ukrainiansd,f | 748 | 54 | 22 | 38 | 145 | 2 | 17 | 27 | 43 | 9 | 3 | 338 | 50 |
Unzhaa | 52 | 3 | 0 | 6 | 6 | 2 | 0 | 4 | 7 | 0 | 0 | 17 | 6 |
Vologdaa | 121 | 1 | 3 | 14 | 10 | 2 | 0 | 1 | 47 | 0 | 2 | 40 | 1 |
a Balanovsky et al., 2008 ; b Mirabal et al., 2009 ; c Kushniarevich et al., 2013 ; d Kushniarevich et al., 2015 ; eBoattini et al., 2013; f Battaglia et al., 2009 ; g Karachanak et al., 2013 ); hSolé-Morata et al., 2015; iKing et al., 2008; j Karlsson et al., 2006 ; k Rebala et al., 2013 ; lMartinez-Cruz et al., 2012; mVarzari et al., 2013; nReguiero et al., 2012.