The ancient Hungarians originated from the Ural region of Russia, and migrated through the Middle-Volga region and the Eastern European steppe into the Carpathian Basin during the ninth century AD. Their Homeland was probably in the southern Trans-Ural region, where the Kushnarenkovo culture was disseminated. In the Cis-Ural region Lomovatovo and Nevolino cultures are archaeologically related to ancient Hungarians. In this study we describe maternal and paternal lineages of 36 individuals from these regions and nine Hungarian Conquest period individuals from today’s Hungary, as well as shallow shotgun genome data from the Trans-Uralic Uyelgi cemetery. We point out the genetic continuity between the three chronological horizons of Uyelgi cemetery, which was a burial place of a rather endogamous population. Using phylogenetic and population genetic analyses we demonstrate the genetic connection between Trans-, Cis-Ural and the Carpathian Basin on various levels. The analyses of this new Uralic dataset fill a gap of population genetic research of Eurasia, and reshape the conclusions previously drawn from tenth to eleventh century ancient mitogenomes and Y-chromosomes from Hungary.

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After 568 AD the Avars settled in the Carpathian Basin and founded the Avar Qaganate that was an important power in Central Europe until the 9th century. Part of the Avar society was probably of Asian origin; however, the localisation of their homeland is hampered by the scarcity of historical and archaeological data. Here, we study mitogenome and Y chromosomal variability of twenty-six individuals, a number of them representing a well-characterised elite group buried at the centre of the Carpathian Basin more than a century after the Avar conquest. The studied group has maternal and paternal genetic affinities to several ancient and modern East-Central Asian populations. The majority of the mitochondrial DNA variability represents Asian haplogroups (C, D, F, M, R, Y and Z). The Y-STR variability of the analysed elite males belongs only to five lineages, three N-Tat with mostly Asian parallels and two Q haplotypes. The homogeneity of the Y chromosomes reveals paternal kinship as a cohesive force in the organisation of the Avar elite strata on both social and territorial level. Our results indicate that the Avar elite arrived in the Carpathian Basin as a group of families, and remained mostly endogamous for several generations after the conquest. The Carpathian Basin in East-Central Europe is generally regarded as the westernmost point of the Eurasian steppe, and as such, its history was often influenced by the movements of nomadic people of eastern origin. After 568 AD, the Avars settled in the Carpathian Basin and founded their empire which was a powerful player in the geopolitical arena of Central and Eastern Europe for a quarter of a millennium 1,2. The hypothesis of the Asian origin of the Avars appeared as early as the 18th century. Since then various research approaches emerged indicating different regions as their home of origin: i.e. Central or East-Central Asia (see SI chapter 1b for explanation of this geographic term). This debate remained unresolved, however a rising number of evidences points towards the latter one 1,2. The history of the Avars is known from external, mainly Byzantine written accounts of diplomatic and historical character focusing on certain events and important people for the Byzantine Empire. As an example, the description of a Byzantine diplomatic mission in 569-570 AD visiting the Western Turkic Qaganate in Central Asia, claimed that their ruler complained about the escape of his subjects, the Avars 2-4. The linguistic data concerning the Avars are limited to a handful of personal names and titles (Qagan, Bayan, Yugurrus, Tarkhan, etc.) mostly of East-Central Asian origin, known from the same Byzantine written accounts.

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 medieval 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 originates 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 analysed 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 genepool. 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 genepool 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.

28After 568 AD the Avars settled in the Carpathian Basin and founded the Avar Qaganate that 29 was an important power in Central Europe until the 9 th century. Part of the Avar society was 30 probably of Asian origin, however the localisation of their homeland is hampered by the 31 scarcity of historical and archaeological data. 33Here, we study mitogenome and Y chromosomal STR variability of twenty-six individuals, a 34 number of them representing a well-characterised elite group buried at the centre of the 35 Carpathian Basin more than a century after the Avar conquest. 36The studied group has maternal and paternal genetic affinities to several ancient and modern 37 East-Central Asian populations. The majority of the mitochondrial DNA variability represents 38 Asian haplogroups (C, D, F, M, R, Y and Z). The Y-STR variability of the analysed elite males 39 belongs only to five lineages, three N-Tat with mostly Asian parallels and two Q haplotypes.40 The homogeneity of the Y chromosomes reveals paternal kinship as a cohesive force in the 41 organisation of the Avar elite strata on both social and territorial level. Our results indicate that 42 the Avar elite arrived in the Carpathian Basin as a group of families, and remained mostly 43 endogamous for several generations after the conquest.44

Here we present 115 whole mitogenomes and 92 Y-chromosomal Short Tandem Repeat (STR) and Single Nucleotide Polymorphism (SNP) profiles from a Hungarian ethnic group, the Székelys (in Romanian: Secuii, in German: Sekler), living in southeast Transylvania (Romania). The Székelys can be traced back to the 12th century in the region, and numerous scientific theories exist as to their origin. We carefully selected sample providers that had local ancestors inhabiting small villages in the area of Odorheiu Secuiesc/Székelyudvarhely in Romania. The results of our research and the reported data signify a qualitative leap compared to previous studies since it presents the first complete mitochondrial DNA sequences and Y-chromosomal profiles of 23 STRs from the region. We evaluated the results with population genetic and phylogenetic methods in the context of the modern and ancient populations that are either geographically or historically related to the Székelys. Our results demonstrate a predominantly local uniparental make-up of the population that also indicates limited admixture with neighboring populations. Phylogenetic analyses confirmed the presumed eastern origin of certain maternal (A, C, D) and paternal (Q, R1a) lineages, and, in some cases, they could also be linked to ancient DNA data from the Migration Period (5th–9th centuries AD) and Hungarian Conquest Period (10th century AD) populations.

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. Here we present the uniparental genetic makeup of the medieval 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 originates from ancient proto-Ob-Ugric people from Western Siberia (6-13th century), pre-Conquest period and subsisting Hungarians from Volga-Ural region (6-14th century) and their neighbours. By examining the Hungarian prehistory's eastern archaeological traits, we also study their genetic composition and origin in an interdisciplinary framework. We analysed 110 deep sequenced mitogenomes and 42 Y-chromosome haplotypes from 18 archaeological sites from 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 genepool. Based on phylogenetics, we demonstrate and model the connections and splits of the studied Volga-Ural and conqueror groups. Early Hungarians and their alliances conquested the Carpathian Basin in the 890s AD. Re-analysis of the Hungarian conquerors' maternal genepool 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 of the area, certain eastern lineages survived for centuries.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.