The History of Slavs Inferred from Complete Mitochondrial Genome Sequences
Marta Mielnik-Sikorska,
Patrycja Daca,
Boris Malyarchuk,
Miroslava Derenko,
Katarzyna Skonieczna,
Maria Perkova,
Tadeusz Dobosz,
Tomasz Grzybowski

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Based on the newest anthropological data it has been suggested that the area between the Oder and Vistula rivers witnessed continuity of human settlement between the Roman period and the early Middle Ages. Indeed, based on morphological features of skeletal materials it has been established that populations of the Przeworsk, Wielbark and Cherniakhovo cultures from the Roman period bear close similarities to the early medieval Western Slavs and not to the medieval Germanic-speaking populations [10], [11]. Furthermore, paleodemographic studies also point to the biological continuity of the populations inhabiting the Oder and Vistula basin in the Roman period and the early medieval Slavic populations of this region [10]. Therefore, anthropological data received thus far make the “allochtonic” hypothesis less plausible, especially in its extreme migrationist form.
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It is also worth noting that some clusters of H5, like H5a could have been of central European origin [14], [24], [27], [30], [31]. Thus, one may assume that phylogenetic reconstruction of H5 together with the information on the geographical distribution of haplotypes belonging to this clade could be helpful in evaluating the origin of Slavs.
Haplogroup H6 is also very interesting because two its branches, H6a and H6b, are characteristic, correspondingly, for European and Central Asian/Near Eastern populations, thus reflecting a long-time separation of Asian and European H6 mtDNA pools [27]. Meanwhile, H6a is relatively frequent in some Slavic populations (such as eastern Slovaks [32]), but its phylogenetic structure on complete mtDNA level is poorly studied yet.
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Results and Discussion
Mitochondrial Control Region Sequences of Slavic Populations
The results of sequence variation and haplogroup assignment of 404 Poles, 157 Ukrainians and 85 Czechs are presented in Table S2.
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Haplogroup C is present with the highest frequency in the populations of central Asia, while its incidence decreases in the areas of southeastern Asia and India. In the populations of Central and Eastern Europe, frequency of haplogroup C is very low [34]. Nevertheless, some subclusters of haplogroup C, like C5c1 clade was reported to occur almost exclusively in Europe [34]. C5c1 haplogroup was previously observed in three Poles [34], four Europeans [46], one Russian [47], one Caucasian and one person of unknown origin [34], [46], [47]. In this study we present an additional haplotype belonging to C5c1 clade, which was found in the Ukrainian mtDNA pool (Figure 4). As the components of the C5c1 haplogroup are virtually absent in Asia and were reported only in the populations of Central Europe, the previous hypothesis by Derenko et al. [34] that the C5c1 clade might be a marker of Siberian ancestry in Central European populations could be further supported by the results of this study. Thus, the presence of C5c1 clade among recent Europeans may reflect their ancient contacts with Asian populations, that could be traced back to the Neolithic period, as the evolutionary age of C5c1 clade was calculated to around 4–9 kya (Table S4).
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The observed close proximity of six Slavic sequences belonging to clades C4a1a, G2a and D5a2a1a1 with samples of Indian and East Asian ancestry seems unexpected. Nevertheless, several reports indicate that these mitochondrial lineages are characterized by a very wide geographical distribution that covers areas of northern, eastern and central Asia [60], [61].
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Subsequent migration through East Asia to Europe might have brought some of these components into Central Europe. Therefore, in respect to haplotypes from D5a2a1a1 clade we could not exclude the possibility that their presence in European population might be associated with relatively recent medieval events that took place in Europe. According to archaeological data, during Middle Ages, Asian populations, including Altaic tribes (Huns, Avars, and Mongols) were engaged in wars on the European continent [4], which could in consequence leave their traces in the form of Asian haplotypes in European populations.
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Conclusions
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These are mainly represented by samples of southern European origin, which further supports the idea of Europe repopulation from southern European refugia after LGM [12], [46], [47]. As expected, we show here that potentially Slavic-specific components of H5 haplogroup are much younger than H5 subclades of southern Europe, as their evolutionary age was calculated to approximately 4 kya. The formation of these clades coincides with the expansion of Central and Eastern European haplogroups U4a2, U5a2a and U5a2b1 [48], [49]. Taken together, this data points to a genetic continuity of several maternal lineages in Central Europe from the times of Bronze and Iron Ages. Interestingly, this picture could be also confirmed by expansion time of Y-chromosome subcluster R1a1a1-M458 [51]. Thus, one may exclude the migrationist assumption that Central European territories were populated by the Slavs only at the very beginning of sixth century, following whole scale depopulation of the northern areas of Central Europe [1]. Indeed, the data presented herein indicates that visible changes of material culture of Central Europe in the fifth century did not result from extensive demographic changes, but were rather accompanied by continuity of some maternal and paternal lineages between Bronze and early Middle Ages.

On the other hand, analysis of entire mitochondrial genomes of Asian and African lineages (A, C, D, G, and L) found in Poles and Ukrainians allows us to reconstruct relatively recent events in the history of Slavs, in terms of their relationships with other ethnic groups. The results presented here provide an additional evidence for the existence of limited maternal gene flow between East Asia and Central Europe. In particular, among Poles we show the presence of A8a1 haplotype, which may reflect the probable medieval migration of the nomadic tribes from Siberia. On the other hand, the existence of haplotypes representing clades C4a1a, G2a and D5a2a1a1 in Polish and Ukrainian populations could further reflect the influx of Asian haplotypes during the Middle Ages wars, in which Altaic tribes were engaged. Furthermore, in the present study we show Ashekanazi-specific L2a1l2a lineage in Polish population, which in addition to K1a1b1a haplotypes previously identified in Poland [23], constitutes further evidence of relatively recent migration of Ashkenazi Jews from Germany to Poland.