Tracing genetic connections of ancient Hungarians to the 6th-14th century populations of the Volga-Ural region

Abstract

Most of the early Hungarian tribes originated from the Volga-Kama and South-Ural regions, where they were composed of a mixed population based on historical, philological and archaeological data. We present here the uniparental genetic makeup of the mediaeval era of these regions that served as a melting pot for ethnic groups with different linguistic and historical backgrounds. Representing diverse cultural contexts, the new genetic data originate from ancient proto-Ob-Ugric people from Western Siberia (6th-13th century), the pre-Conquest period and subsisting Hungarians from the Volga-Ural region (6th-14th century) and their neighbours. By examining the eastern archaeology traits of Hungarian prehistory, we also study their genetic composition and origin in an interdisciplinary framework. We analyzed 110 deep-sequenced mitogenomes and 42 Y-chromosome haplotypes from 18 archaeological sites in Russia. The results support the studied groups' genetic relationships regardless of geographical distances, suggesting large-scale mobility. We detected long-lasting genetic connections between the sites representing the Kushnarenkovo and Chiyalik cultures and the Carpathian Basin Hungarians and confirmed the Uralic transmission of several East Eurasian uniparental lineages in their gene pool. Based on phylogenetics, we demonstrate and model the connections and splits of the studied Volga-Ural and conqueror groups. Early Hungarians and their alliances conquered the Carpathian Basin around 890 AD. Re-analysis of the Hungarian conquerors' maternal gene pool reveals numerous surviving maternal relationships in both sexes; therefore, we conclude that men and women came to the Carpathian Basin together, and although they were subsequently genetically fused into the local population, certain eastern lineages survived for centuries.

Figure 1

The supposed migration route of the early Hungarians (arrows) and the regions that could be linked to them (uppercase letters). The sites were grouped according to archaeological and chronological aspects; the formed groups are marked with white outlines. The investigated sites and the groups formed from them: Bolshie Tigani [1]; Novinki group: Novinki [2], Mulovka [3], Brusyany [4], Lebyazhinka [5], Malaya Ryazan [6], Shilovka [7]; Chiyalik group: Gulyukovo [8], Novo Hozyatovo [9], Gornovo [10]; Tankeevka [11]; Bustanaevo [12]; Proto-Ob-Ugric group: Vikulovo [13], Barshov Gorodok [14], Ivanov Mis [15], Panovo [16], Ust-Tara [17]; Uyelgi + Karanayevo group: Karanayevo [18], Uyelgi [19]; Cis-Ural group: Bayanovo [20], Brody [21], Bartim [22], Sukhoy Log [23] (source of 19–23: Csáky et al. 2020 (22)). Source of the map: Qgis v3.16.0 Topographic WMS-by terrestris (https://ows.terrestris.de/osm/service?). Modifications were made in Adobe Photoshop CS6 and Adobe Photoshop 2020.

Figure 2

The mitochondrial (bar graph) and Y-chromosome (pie charts) haplogroup compositions of the analyzed groups. In the case of the Hungarian conquerors, most of the Y-chromosomal data are known from the KL-IV group (34,35). Owing to the underrepresentation of KL-V and KL-VI groups in the Y-chromosomal dataset, the conquerors were merged into one group in this figure.

Figure 3

Median-joining network of N1a1 Y-chromosomal haplogroup based on 17 STRs. The studied Y lineages from Uyelgi + Karanayevo (Uyelgi and Karanayevo sites) and Chiyalik (Gornovo) groups are identical or closely related to each other and Bashkirs from Perm, Strelibashevsy, Burzyansky and Western Orenburg regions, as well as Khantys, Southern and Northern Mansis, modern-day Hungarians, Tatars and Ukrainians. One sample from the Chiyalik group (Gulyukovo site, GU6) is on a subbranch, mainly composed of Bashkirian Maris. The other sample from this site (GU9) is on another subbranch, which contains Tatars and Russians as well. For information on the groups and STR markers, see Supplementary Material, Table S10.

Figure 4

PCA plot based on mitochondrial haplogroup frequencies of ancient and present-day Eurasian and Near-Eastern populations. The variances presented on the first two components: PC1:16.4%; PC2: 7.1%. For abbreviations and information, see Supplementary Material, Table S3. PC3 with a variance of 6.3% is presented in Supplementary Material, Figure S36.

Figure 5

Shared mtDNA subhaplogroups between the studied groups (under the diagonal) and results of F_ST analysis (over the diagonal). F_ST values in bold indicate a significant genetic distance. In the case of phylogenetic relationships, the more distal but unambiguous relationships are indicated by italics, the normal letters marking close relationships, and letters in bold showing haplotype identity between detected mitogenomes.

Figure 6

Mitochondrial haplogroup N1a1a1a1a phylogenetic tree with divergence dates, made with BEAST software. This tree mainly consists of samples linked to Hungarian prehistory, including both database and data of this study (in bold). Divergence date estimations correlate with presumptions of group split times based on historical and archaeological data. The majority of the group divergence dates are 600–800 AD, pointing to rapid population movements in this period. For the complete tree, see Supplementary Material, Extended Figure 3.