Genetic characterization and meta-analysis of the population of the Northern Black Sea region in the 1st millennium CE based on ancient DNA data

Over the past two decades, the introduction of whole-genome sequencing analysis of ancient DNA has led to a breakthrough in archaeogenetic research, significantly expanding our understanding of human genetic history. In this context the Northern Black Sea region during first millennium CE (1–1,000 CE) is of particular relevance, as it remained a hub of intense cultural exchange and migration. Despite its historical importance, ancient genomic data from this period remains scarce, and a comprehensive synthesis of existing findings is lacking. This study presents a systematic review and meta-analysis of published whole-genome sequencing data from 48 ancient samples associated with key archaeological cultures of the region: Late Scythian, Sarmatian, Alan, Bulgar, Saltovo-Mayaki and Chernyakhov. Through the systematization of data, we trace genetic continuity at Late Scythian and Alanian sites relative to preceding populations. Episodes of large-scale migration and population replacement have been documented, most clearly evident in the Sarmatian expansion of the 1st–4th centuries CE, with genetic traces extending from the Urals to the Carpathians. Based on limited evidence, genetic continuity has been identified between representatives of the Chernyakhov culture and early Slavic groups. Through our meta-analysis, we further detect intercultural connections between the Alans, Bulgars, and bearers of the Saltovo-Mayaki culture, whose genetic structure reveals the influence of Caucasian and East Eurasian components. Collectively, these findings underscore the complex genetic landscape of the region, shaped by successive migration waves and multifaceted intercultural contacts. We conclude by outlining key unresolved questions and future directions for archaeogenetic research in the Northern Black Sea region during the first millennium CE.

1. Afanasyev G.E., Ven’ Sh., Tun S., Van L., Vei L., Dobrovolskaya M.V., Korobov D.S., Reshetova I.K., Li Kh. Khazar Confederates in the Don region. In: Natural Scientific Methods and the Paradigm of Modern Archaeology. Moscow: IA RAN, 2015a;146-154 (in Russian)

2. Afanasyev G.E., Dobrovolskaya M.V., Korobov D.S., Reshetova I.K. New archaeological, anthropological and genetic aspects in the study of the Alans from the Don region. Kratkiye Soobshcheniya Instituta Arkheologii = Brief Communications of the Institute of Archaeology. 2015b;(237):64-79 (in Russian)

3. Alekseeva E.M. Greek Colonization of the Northwestern Caucasus. Moscow, 1991 (in Russian)

4. Alexander D.H., Novembre J., Lange K. Fast model-based estimation of ancestry in unrelated individuals. Genome Res. 2009;19(9):1655-1664. doi 10.1101/gr.094052.109

5. Allentoft M.E., Sikora M., Sjögren K.-G., Rasmussen S., Rasmussen M., Stenderup J., Damgaard P.B., … Sicheritz-Pontén T., Brunak S., Nielsen R., Kristiansen K., Willerslev E. Population genomics of Bronze Age Eurasia. Nature. 2015;522(7555):167-172. doi 10.1038/nature14507

6. Allentoft M.E., Sikora M., Refoyo-Martínez A., Irving-Pease E.K., Fischer A., Barrie W., Ingason A., ... Delaneau O., Werge T., Racimo F., Kristiansen K., Willerslev E. Population genomics of post-glacial western Eurasia. Nature. 2024;625(7994):301-311. doi 10.1038/s41586-023-06865-0

7. Andreeva T.V., Soshkina A.D., Gusev F.E., Malyarchuk A.B., Dotsenko G.S., Dudko N.A., Plotnikova M.Yu., … Kantorovich A.R., Maslov V.E., Barinov D.G., Dobrovolskaya M.V., Rogaev E.I. Genetic history of Scythia. Sci Adv. 2025:11(30):eads8179. doi 10.1126/sciadv.ads8179

8. Berezina N.Y., Buzhilova A.P., Reshetova I.K. New craniological materials revisited anthropological background of medieval Alans. Vestnik MGU. Seriya XXIII. Antropologiya = Lomonosov J Anthropol. 2012;(4):18-36 (in Russian)

9. Bubenok O.B. To the Question about the Quantity of Tribes of the Nomads of North Black Sea Region in the late 5–frst half of 7 century AD. Drinovs’kij Zbirnik. Kharkiv; Sofiya. 2014, V. IX;29-35

10. Comas D., Plaza S., Wells R.S., Yuldaseva N., Lao O., Calafell F., Bertranpetit J. Admixture, migrations, and dispersals in Central Asia: evidence from maternal DNA lineages. Eur J Hum Genet. 2004; 12(6):495-504. doi 10.1038/sj.ejhg.5201160

11. Damgaard P.B., Marchi N., Rasmussen S., Peyrot M., Renaud G., Korneliussen T., Moreno-Mayar J.V., … Nielsen R., Sikora M., Heyer E., Kristiansen K., Willerslev E. 137 ancient human genomes from across the Eurasian steppes. Nature. 2018;557(7705):369-374. doi 10.1038/s41586-018-0094-2

12. Dashevskaya O.D. The Belyaus Necropolis. Simferopol, 2014 (in Russian)

13. Derenko M., Malyarchuk B., Grzybowski T., Denisova G., Rogalla U., Perkova M., Dambueva I., Zakharov I. Origin and post-glacial dispersal of mitochondrial DNA haplogroups C and D in northern Asia. PloS One. 2010:5(12):e15214. doi 10.1371/journal.pone.0015214

14. Derenko M., Malyarchuk B., Denisova G., Perkova M., Rogalla U., Grzybowski T., Khusnutdinova E., Dambueva I., Zakharov I. Complete mitochondrial DNA analysis of eastern Eurasian haplogroups rarely found in populations of Northern Asia and Eastern Europe. PLoS One. 2012;7(2):e32179. doi 10.1371/journal.pone.0032179

15. ENA Browser, 2025. Availabe online: https://www.ebi.ac.uk/ena/browser/home

16. Gelabert P., Schmidt R.W., Fernandes D.M., Karsten J.K., Harper T.K., Madden G.D., Ledogar S.H., Sokhatsky M., Oota H., Kennett D.J., Pinhasi R. Genomes from Verteba cave suggest diversity within the Trypillians in Ukraine. Sci Rep. 2022;12(1):7242. doi 10.1038/s41598-022-11117-8

17. Haak W., Lazaridis I., Patterson N., Rohland N., Mallick S., Llamas B., Brandt G., … Brown D., Anthony D., Cooper A., Alt K.W., Reich D. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature. 2015;522(7555):207-211. doi 10.1038/nature14317

18. Harney É., Patterson N., Reich D., Wakeley J. Assessing the performance of qpAdm: a statistical tool for studying population admixture. Genetics. 2021;217(4):iyaa045. doi 10.1093/genetics/iyaa045

19. Järve M., Saag L., Scheib C.L., Pathak A.K., Montinaro F., Pagani L., Flores R., … Metspalu M., Savelev N., Kriiska A., Kivisild T., Villems R. Shifts in the genetic landscape of the Western Eurasian Steppe associated with the beginning and end of the Scythian dominance. Curr Biol. 2019;29(14):2430-2441.e10. doi 10.1016/j.cub.2019.06.019

20. Jones E.R., Zarina G., Moiseyev V., Lightfoot E., Nigst P.R., Manica A., Pinhasi R., Bradley D.G. The Neolithic transition in the Baltic was not driven by admixture with early European farmers. Curr Biol. 2017;27(4):576-582. doi 10.1016/j.cub.2016.12.060

21. Juras A., Krzewińska M., Nikitin A.G., Ehler E., Chyleński M., Łukasik S., Krenz-Niedbała M., Sinika V., Piontek J., Ivanova S., Dabert M., Götherström A. Diverse origin of mitochondrial lineages in Iron Age Black Sea Scythians. Sci Rep. 2017;7(1):43950. doi 10.1038/srep43950

22. Kazanski M.M. Radagais and the End of the Chernyakhov Culture (Radagaïs et la fin de la civilisation de Černjahov). OIUM. Kyiv; Lutsk. Vol. 1. 2011;1:22-32 (in Russian)

23. Kazanski M.M. the antiquities of the steppe nomads of the Post-Hunnic period (mid-fifth to mid-sixth century in Eastern Europe. Materialy po Arkheologii, Istorii i Etnografii Tavrii = Materials in Archaeology, History and Ethnography of Tauria. 2020;25:90-167 (in Russian)

24. Korobov D.S. The Alans of the North Caucasus: Ethnos, Archaeology, Paleogenetics. Saint Petersburg, 2019 (in Russian)

25. Koryakova L. Europe to Asia. In: The Oxford Handbook of the European Iron Age. Oxford University Press, 2018;575-618. doi 10.1093/oxfordhb/9780199696826.013.29

26. Koshelenko G.A., Kruglikova I.T., Dolgorukov V.S. Ancient States of the Northern Black Sea Region. Moscow, 1984 (in Russian)

27. Kovács P. Fontes Sarmatarum in Hungaria Habitantium – A Magyarországi Szarmatákra Vonatkozó Antik Forrásaink. Budapest: Magyarságkutató Intézet, 2023

28. Krzewińska M., Kılınç G.M., Juras A., Koptekin D., Chyleński M., Nikitin A.G., Shcherbakov N., … Dalén L., Sinika V., Jakobsson M., Storå J., Götherström A. Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western Iron Age nomads. Sci Adv. 2018;4(10):eaat4457. doi 10.1126/sciadv.aat4457

29. Lazaridis I., Nadel D., Rollefson G., Merrett D.C., Rohland N., Mallick S., Fernandes D., … Yengo L., Hovhannisyan N.A., Patterson N., Pinhasi R., Reich D. Genomic insights into the origin of farming in the ancient Near East. Nature. 2016;536(7617):419-424. doi 10.1038/nature19310

30. Lazaridis I., Alpaslan-Roodenberg S., Acar A., Açıkkol A., Agelarakis A., Aghikyan L., Akyüz U., … Çavuşoğlu R., Rohland N., Pinhasi R., Reich D., Davtyan R. The genetic history of the Southern Arc: a bridge between West Asia and Europe. Science. 2022;377(6609): eabm4247. doi 10.1126/science.abm4247

31. Malashev V.Yu., Krivosheev M.V. Revisiting the issue of attribution of some catacomb monuments groups from the south of Eastern Europe in the late 3rd–early 5th centuries. Nizhnevolzhskii Archaeologicheskii Vestnik = Lower Volga Archaeological Bulletin. 2024; 23(3):52-78. doi 10.15688/nav.jvolsu.2024.3.4 (in Russian)

32. Mallick S., Micco A., Mah M., Ringbauer H., Lazaridis I., Olalde I., Patterson N., Reich D. The Allen Ancient DNA Resource (AADR) a curated compendium of ancient human genomes. Sci Data. 2024; 11(1):182. doi 10.1038/s41597-024-03031-7

33. Maróti Z., Neparáczki E., Schütz O., Maár K., Varga G.I.B., Kovács B., Kalmár T., … Gulyás B., Kovacsóczy B.N., Gál S.S., Tomka P., Török T. The genetic origin of Huns, Avars, and conquering Hungarians. Curr Biol. 2022;32(13):2858-2870.e7. doi 10.1016/j.cub.2022.04.093

34. Mathieson I., Alpaslan-Roodenberg S., Posth C., Szécsényi-Nagy A., Rohland N., Mallick S., Olalde I., … Borić D., Bonsall C., Krause J., Pinhasi R., Reich D. The genomic history of southeastern Europe. Nature. 2018;555(7695):197-203. doi 10.1038/nature25778

35. Mattila T.M., Svensson E.M., Juras A., Günther T., Kashuba N., Ala-Hulkko T., Chyleński M., … Netea M.G., Nikitin A.G., Persson P., Malmström H., Jakobsson M. Genetic continuity, isolation, and gene flow in Stone Age Central and Eastern Europe. Commun Biol. 2023; 6(1):793. doi 10.1038/s42003-023-05131-3

36. Matveev S.V. Barbarized Romans or Romanized Barbarians in the Chernyakhov Community of the Prut-Dniester Interfluve. In: Eastern Europe in Antiquity and the Middle Ages: Ancient and Medieval Communities. Moscow, 2017;147-151 (in Russian)

37. McVean G. A genealogical interpretation of principal components analysis. PLoS Genet. 2009;5(10):e1000686. doi 10.1371/journal.pgen.1000686

38. Mordvintseva V. The Sarmatians: the creation of archaeological evidence. Oxford J. Archaeol. 2013;32(2):203-219. doi 10.1111/ojoa.12010

39. Nikitin A.G., Lazaridis I., Patterson N., Ivanova S., Videiko M., Dergachev V., Kotova N., … Workman J.N., Zalzala F., Mallick S., Rohland N., Reich D. A genomic history of the North Pontic Region from the Neolithic to the Bronze Age. Nature. 2025;639(8053):124-131. doi 10.1038/s41586-024-08372-2

40. Onishchuk Ja.I. Relationship of the populations of the Velbar, Sarmatian, and Chernyakhov cultures in the territory of Western Podolia in the second quarter of the 1st millennium AD. Zhurnal Belorusskogo Gosudarstvennogo Universiteta. Istoriya = Journal of the Belarusian State University. History. 2018;3:120-129 (in Russian)

41. Patterson N., Price A.L., Reich D. Population structure and eigen-analysis. PLoS Genet. 2006;2(12):e190. doi 10.1371/journal.pgen.0020190

42. Patterson N., Moorjani P., Luo Y., Mallick S., Rohland N., Zhan Y., Genschoreck T., Webster T., Reich D. Ancient admixture in human history. Genetics. 2012;192(3):1065-1093. doi 10.1534/genetics.112.145037

43. Patterson N., Isakov M., Booth T., Büster L., Fischer C.E., Olalde I., Ringbauer H., ... Lillie M., Rohland N., Pinhasi R., Armit I., Reich D. Large-scale migration into Britain during the Middle to Late Bronze Age. Nature. 2021;601(7894):588-594. doi 10.1038/s41586-021-04287-4

44. Penske S., Rohrlach A.B., Childebayeva A., Gnecchi-Ruscone G., Schmid C., Spyrou M.A., Neumann G.U., ... Ringbauer H., Stockhammer P.W., Hansen S., Krause J., Haak W. Early contact between late farming and pastoralist societies in southeastern Europe. Nature. 2023;620(7973):358-365. doi 10.1038/s41586-023-06334-8

45. Perеvalov S.M. Alans: a Mirage of Nomadic Empire. Vestnik Vladikavkazskogo Nauchnogo Tsentra = Vestnik of Vladikavkaz Scientific Centre. 2014;14(2):2-9 (in Russian)

46. Pletneva S.A. Nomads of the South Russian Steppes in the Middle Ages (4th–13th Centuries). Voronezh, 2003 (in Russian)

47. Post H., Németh E., Klima L., Flores R., Fehér T., Türk A., Székely G., … Khusnutdinova E.K., Metspalu E., Villems R., Tambets K., Rootsi S. Y-chromosomal connection between Hungarians and geographically distant populations of the Ural Mountain region and West Siberia. Sci Rep. 2019;9(1):7786. doi 10.1038/s41598-019-44272-6

48. Posth C., Yu H., Ghalichi A., Rougier H., Crevecoeur I., Huang Y., Ringbauer H., ... Terberger T., Caramelli D., Altena E., Haak W., Krause J. Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers. Nature. 2023;615(7950):117-126. doi 10.1038/s41586-023-05726-0

49. Reshetova I.K. New anthropological materials of the Saltovo-Mayaki culture from the Verkhny Saltov-IV burial ground. Rossijskaya Archeologiya = Russian Archaeology. 2012;(3):129-136 (in Russian) Ringbauer H., Huang Y., Akbari A., Mallick S., Olalde I., Patterson N., Reich D. Accurate detection of identity-by-descent segments in human ancient DNA. Nat Genet. 2024;56(1):143-151. doi 10.1038/s41588-023-01582-w

50. Rozhdestvenskikh E.V., Andreeva T.V., Malyarchuk A.B., Adrianova I.Yu., Khodyreva D.S., Evteev A.A., Buzhilova A.P., Rogaev E.I. Mitogenomic analysis of a representative of the Chernyakhov culture in the Middle Dniester and their genetic relationship with the Slavs in the context of paleoanthropological data. Vavilovskii Zhurnal Genetikii Selektsii = Vavilov J Genet Breed. 2025;29(5):722-731. doi 10.18699/vjgb-25-79

51. Saag L., Utevska O., Zadnikov S., Shramko I., Gorbenko K., Bandrivskyi M., Pavliv D., … Barrington C., Gilardet A., Macleod R., Skoglund P., Thomas M.G. North Pontic crossroads: mobility in Ukraine from the Bronze Age to the early modern period. Sci Adv. 2025;11(2):eadr0695. doi 10.1126/sciadv.adr0695

52. Schütz O., Maróti Z., Tihanyi B., Kiss A.P., Nyerki E., Gînguță A., Kiss P., … Soficaru A.D., Spekker O., Varga S., Neparáczki E., Török T. Unveiling the origins and genetic makeup of the “forgotten people”: a study of the Sarmatian-period population in the Carpathian Basin. Cell. 2025;188(15):4074-4090.e11. doi 10.1016/j.cell.2025.05.009

53. Sedov V.V. The Origin and Early History of the Slavs. Moscow, 1979 (in Russian)

54. Sharko F.S., Boulygina E.S., Tsygankova S.V., Slobodova N.V., Rastorguev S.M., Krasivskaya A.A., Belinsky A.B., … Shvedchikova T.Yu., Dobrovolskaya M.V., Reshetova I.K., Korobov D.S., Nedoluzhko A.V. Koban culture genome-wide and archeological data open the bridge between Bronze and Iron Ages in the North Caucasus. Eur J Hum Genet. 2024;32(11):1483-1491. doi 10.1038/s41431-023-01524-4

55. Shushunova E.V., Yartsev S.V. The Ancestor Cult of the Ancient Population of the Northern Black Sea Region in the 1st Century BC – 4th Century AD and Features of its Further Development in the Way of Transformation from Paganism to Christianity. Simferopol, 2024 (in Russian)

56. Wang C.C., Reinhold S., Kalmykov A., Wissgott A., Brandt G., Jeong C., Cheronet O., ... Belinskij A.B., Reich D., Hansen S., Krause J., Haak W. Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions. Nat Commun. 2019;10(1):590. doi 10.1038/s41467-018-08220-8

57. YTree, 2025. Availabe online: https://www.yfull.com/tree/