The Scythians: Who Were They?

Figure 5: Median network of U5 haplotypes.

Representatives of the U5 haplogroup account for one third of the mt haplotypes identified in analyzed Scythians. These ultimately relate to West Eurasian hunter-gatherers, whose descendants extended throughout the European subcontinent and into East Eurasia from the Mesolithic to the Bronze Age. We have combined the known diversity of the prehistoric U5 lineage from different sources19,45,46,47,52,53,54,55,56 into a median network, which allowed for identification of candidate source populations contributing to the U5a diversity in NPR Scythians. Although the network approach is still limited by the small number of available samples, our results indicate that the source populations with the most closely related haplotypes are the Bronze Age Srubnaya (SRU) (1900-1200 BCE) and the earlier Yamnaya (YAM) (3300-2700 BCE) from the Ponto-Caspian region and Bronze Age populations from the Altai Mountains, such as Karasuk (1500-800 BCE). Diverse U5a-carrying populations of the steppe such as Yamnaya, Srubnaya and Scythians shared the nomadic lifestyle, with the economic foundations transforming from wild game exploitation in the Mesolithic to pastoralic animal husbandry in the Bronze Age. Furthermore, both Yamnaya and Srubnaya were part of the Kurgan culture phenomenon57 with the Scythian cultural horizon being the most representative for the kurgan building tradition.

The second group of identified mt genomes in the NPR Scythians is comprised of lineages that suggest associations of the NPR Scythians with the Neolithic European farming groups. Although the mt lineage composition of analyzed Scythians significantly differs from that seen in NEN and LBK groups (FST = 0.06, FST = 0.04, respectively, p > 0.05), particular lineages such as J1c2, T2b, H/H5, H, W and N1b1a ultimately go back to the earliest European farmers. Lineages of J1c, H5 and T2b belong to the Neolithic farming package of mtDNA haplogroups which have been found in most Neolithic and Bronze Age European populations19,44,47,55,58. Noteworthy, individuals belonging to the N1b1a have been found in the Neolithic Anatolia19,59, but the lineage has not appeared in any other Eurasian Neolithic, Bronze or Iron Age populations to date. Therefore, the presence of N1b1a in Scythians could be attributed to population migrations along the southern boundaries of the Ponto-Caspian region60.

Mitochondrial lineages T2b, W3a and J2b1a6 have been identified in representatives of the Bronze Age groups such as Srubnaya (T2b, J2b1a, H5)19, Sintashta (J2b1a, J1c)55, Yamnaya (W3a and T2b)47,61 and Mezhovskaya (J2b1a)55 along with East European Eneolithic Trypillian culture (T2b)62, suggesting genetic continuity of these lineages from at least the Bronze Age or even Neolithic times in NPR region. This is further supported by published data19 which showed that almost one fifth of the genetic makeup of the Late Bronze Age Srubnaya people of the Ponto-Caspian region is of the Early European Farmer or Anatolian Neolithic ancestry possibly resulting from the admixture of populations related to Early European Farmers and Yamnaya. Thus, if the NPR Scythians are the descendants of populations related to Srubnaya, the origin of the identified farming lineages would likely be within the steppe/forest-steppe region between the NPR and southern Ural. According to previous genomic studies47,55 the CWC people are likely to have arisen from Yamnaya background. Thus, genetic affinities of the NPR Scythians to the Yamnaya people might also explain their close genetic similarity to the CWC reflected in PCA and FST results. Close genetic relations of the NPR Scythians and Srubnaya are supported by the FST analyses revealing no significant differences between these two populations (Figs 3 and 4, Supplementary Table S5). AMOVA further corroborated these results showing that the combination of Scythians and Srubnaya results in the lowest genetic variability among populations in a group and the highest variability between groups (Supplementary Table S6). Our results support the archaeological interpretations concerning the origin of the Scythians. One of the versions of this theory suggests that the Srubnaya people have migrated in several waves from the Volga-Ural region to the NPR during the second half of the second millennium BCE where their descendants gave rise to the Scythians around the 7th century BCE63.



The Pazyryks from Mongolia and Altai were separated from the Scythians from present day Moldova and Ukraine (SCU) in the PC analysis (Figure 2) and they clustered with Central and East Asian populations (Juras et al. 2017). West Eurasian haplogroups U5 and H are dominant in the mtDNA haplogroup profile of the Scythians from Moldova and Ukraine, while the Pazyryks from Mongolia and Altai have more East Eurasian mtDNA lineages, such as A, D4j2 and F1b, compared to their SCU counterparts. East Eurasian mtDNA lineages increase from 26.3% in the Scythians (SCU) to 46.7% in the Pazyryks from Mongolia. Since haplogroup U5 accounts for 31.6% of the mtDNA haplotypes of the Scythians studied, they are genetically related to West Eurasian hunter-gatherers, whose descendants extended throughout the European subcontinent.

The third group of lineages identified in the NPR Scythians is derived from East Eurasian ancestry. Since, to our knowledge, there is no evidence of agricultural subsistence in East Eurasian Scythians, these lineages should be considered to be genetic components associated with nomadic populations. Mitochondrial haplogroups such as A, D and F have already been found in samples from the Mesolithic, Neolithic, Bronze and Iron Age southern Siberia and the Altai7,14,15,16,18,55,64,65,66,67. Notably, haplogroup M10 found among Scythians from Glinoe, is present not only in far East Asia but also in modern populations of the Altai68. Among ancient populations, the M10 lineages have been found in Chinese specimens from southern Xinjiang48 (8th -1st century BCE) and Xiongnu69 (3rd century BCE). Furthermore, haplogroup M have been identified in Pleistocene individuals from western Europe but it is thought that these lineages disappeared due to bottleneck effect in the Last Glacial Maximum56. Haplogroup H8c, identified in the NPR Scythians likely belongs to East Eurasian lineages as well. Sequence analysis of 830 modern Eurasian mt genomes suggested a distinct phylogeographc history for H8, with a clustering of Near Eastern and Central Asian haplotypes of H8 and a pronounced presence of carriers of H8c in the Altai region70. It was further hypothesized that the distribution of this lineage could have been facilitated by nomadic migrations along the NPR coast70. Although our PCA analysis showed Altai Pazyryk (SCA) to be distant from the NPR Scythians (SCU) (Fig. 2), it must be emphasized that the presence of haplogroups A and H8c in the analyzed population connects NPR Scythians to the Altai and identifies this region as a possible source of these East Eurasian mt lineages. The only Scythian mt genome from southern Urals published thus far also belonged to an East Eurasian lineage, G2a19. The Y chromosomal lineage (R1a1a1b2a2a) reconstructed for this individual was supposedly characteristic for members of the Srubnaya culture19 which additionally supports our conclusions concerning close genetic links between Scythians and people related to the late Bronze Age Srubnaya.

Scythians from Rostov-on-Don as well as Pazyryks from Altai and Inner Mongolia were carriers of mixed east and west Eurasian lineages, with the dominant presence of the latter at 62.5% and 53.3%, respectively7,13,14,15,16,17,18. Mitochondrial haplogroup analyses of the NPR Scythians from this study and those from Rostov-on-Don and Pazyryks from Altai and Inner Mongolia, reveal that, for the most part, the same lineages are found in all three groups and are often singularly represented in each group. Noteworthy, comparing the frequencies of east and west Eurasian haplogroups in all three groups of the Scythian horizon, an east-west mtDNA lineage cline is visible, for east Eurasian lineages going west-east is from 26.3% (in present study) through 37.5% (in Scythians from Rostov-on-Don) to 46.7% (in Pazyryks) with the opposite trend for west Eurasian lineages.


Scientific Reports volume 7, Article number: 43950 (2017)