A genetic history of the Balkans from Roman frontier to Slavic migrations

Iñigo Olalde¹, Pablo Carrión², Ilija Mikić³, Nadin Rohland⁴, Swapan Mallick⁵, Iosif Lazaridis⁶, Matthew Mah⁷, Miomir Korać³, Snežana Golubović³, Sofija Petković³, Nataša Miladinović-Radmilović³, Dragana Vulović³, Timka Alihodžić⁸, Abigail Ash⁹, Miriam Baeta¹⁰, Juraj Bartík¹¹, Željka Bedić¹², Maja Bilić¹³, Clive Bonsall¹⁴, Maja Bunčić¹⁵, Domagoj Bužanić¹⁶, Mario Carić¹², Lea Čataj¹⁷, Mirna Cvetko¹⁶, Ivan Drnić¹⁵, Anita Dugonjić¹⁵, Ana Đukić¹⁵, Ksenija Đukić¹⁸, Zdeněk Farkaš¹¹, Pavol Jelínek¹¹, Marija Jovanovic¹⁹, Iva Kaić¹⁶, Hrvoje Kalafatić²⁰, Marijana Krmpotić²¹, Siniša Krznar²⁰, Tino Leleković²², Marian M de Pancorbo¹⁰, Vinka Matijević¹⁶, Branka Milošević Zakić²³, Anna J Osterholtz²⁴, Julianne M Paige²⁵, Dinko Tresić Pavičić²⁶, Zrinka Premužić²⁷, Petra Rajić Šikanjić¹², Anita Rapan Papeša²⁸, Lujana Paraman²⁹, Mirjana Sanader¹⁶, Ivana Radovanović³⁰, Mirjana Roksandic³¹, Alena Šefčáková³², Sofia Stefanović³³, Maria Teschler-Nicola³⁴, Domagoj Tončinić¹⁶, Brina Zagorc³⁵, Kim Callan⁵, Francesca Candilio³⁶, Olivia Cheronet³⁵, Daniel Fernandes³⁷, Aisling Kearns⁴, Ann Marie Lawson⁵, Kirsten Mandl³⁵, Anna Wagner³⁵, Fatma Zalzala⁵, Anna Zettl³⁵, Željko Tomanović³⁸, Dušan Keckarević³⁹, Mario Novak¹², Kyle Harper⁴⁰, Michael McCormick⁴¹, Ron Pinhasi⁴², Miodrag Grbić⁴³, Carles Lalueza-Fox⁴⁴, David Reich⁴⁵

Affiliations

¹ BIOMICs Research Group, Department of Zoology and Animal Cell Biology, University of the Basque Country UPV/EHU, Vitoria-Gasteiz, Spain; Ikerbasque-Basque Foundation of Science, Bilbao, Spain; Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA; Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, Spain. Electronic address: inigo.olalde@ehu.eus.
² Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, Spain.
³ Institute of Archaeology, Belgrade, Serbia.
⁴ Department of Genetics, Harvard Medical School, Boston, MA, USA.
⁵ Department of Genetics, Harvard Medical School, Boston, MA, USA; Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA.
⁶ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA; Department of Genetics, Harvard Medical School, Boston, MA, USA.
⁷ Department of Genetics, Harvard Medical School, Boston, MA, USA; Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA.
⁸ Archaeological Museum Zadar, Zadar, Croatia.
⁹ Department of Archaeology, Durham University, Durham, UK.
¹⁰ BIOMICs Research Group, Department of Zoology and Animal Cell Biology, University of the Basque Country UPV/EHU, Vitoria-Gasteiz, Spain.
¹¹ Slovak National Museum-Archaeological Museum, Bratislava, Slovak Republic.
¹² Centre for Applied Bioanthropology, Institute for Anthropological Research, Zagreb, Croatia.
¹³ Palisada Ltd., Split, Croatia.
¹⁴ School of History, Classics and Archaeology, University of Edinburgh, Edinburgh, UK.
¹⁵ Archaeological Museum in Zagreb, Zagreb, Croatia.
¹⁶ Faculty of Humanities and Social Sciences, University of Zagreb, Zagreb, Croatia.
¹⁷ Division for Archaeological Heritage, Croatian Conservation Institute, Zagreb, Croatia.
¹⁸ Center of Bone Biology, Faculty of Medicine, University of Belgrade, Belgrade, Serbia.
¹⁹ Museum of Vojvodina, Novi Sad, Serbia.
²⁰ Institute of Archaeology, Zagreb, Croatia.
²¹ Department for Archaeology, Croatian Conservation Institute, Zagreb, Croatia.
²² Archaeology Division, Croatian Academy of Sciences and Arts, Zagreb, Croatia.
²³ Museum of Croatian Archaeological Monuments, Split, Croatia.
²⁴ Department of Anthropology and Middle Eastern Cultures, Mississippi State University, Starkville, MS, USA.
²⁵ Department of Anthropology, University of Nevada, Las Vegas, NV, USA.
²⁶ Kaducej Ltd., Split, Croatia.
²⁷ Independent Researcher, 10000 Zagreb, Croatia.
²⁸ Vinkovci Municipal Museum, Vinkovci, Croatia.
²⁹ Trogir Town Museum, Trogir, Croatia.
³⁰ Department of Anthropology, University of Kansas, Lawrence, KS, USA.
³¹ Department of Anthropology, University of Winnipeg, Winnipeg, MB, Canada.
³² Department of Anthropology, Slovak National Museum-Natural History Museum, Bratislava, Slovak Republic.
³³ Laboratory for Bioarchaeology, Faculty of Philosophy, University of Belgrade, Belgrade, Serbia.
³⁴ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria; Department of Anthropology, Natural History Museum Vienna, Vienna, Austria.
³⁵ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria.
³⁶ Servizio di Bioarcheologia, Museo delle Civiltà, Rome, Italy.
³⁷ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria; Research Centre for Anthropology and Health (CIAS), Department of Life Sciences, University of Coimbra, Coimbra, Portugal.
³⁸ Faculty of Biology, University of Belgrade, Belgrade, Serbia; Serbian Academy of Sciences and Arts, Belgrade, Serbia.
³⁹ Faculty of Biology, University of Belgrade, Belgrade, Serbia.
⁴⁰ Department of Classics and Letters, University of Oklahoma, Norman, OK, USA; Santa Fe Institute, Santa Fe, NM, USA.
⁴¹ Department of History, Harvard University, Cambridge, MA, USA; Max Planck-Harvard Research Center for the Archaeoscience of the Ancient Mediterranean, Harvard University, Cambridge, MA, USA.
⁴² Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria; Human Evolution and Archaeological Sciences, University of Vienna, Vienna, Austria.
⁴³ Faculty of Biology, University of Belgrade, Belgrade, Serbia; Department of Biology, University of Western Ontario, London, ON, Canada; Department of Agriculture and Food, Universidad de La Rioja, Logroño, Spain.
⁴⁴ Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, Spain; Museu de Ciències Naturals de Barcelona, Barcelona, Spain. Electronic address: carles.lalueza.fox@gmail.com.
⁴⁵ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA; Department of Genetics, Harvard Medical School, Boston, MA, USA; Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA. Electronic address: reich@genetics.med.harvard.edu.

PMID: 38065079
PMCID: PMC10752003
DOI: 10.1016/j.cell.2023.10.018

A genetic history of the Balkans from Roman frontier to Slavic migrations

Iñigo Olalde et al. Cell. 2023.

. 2023 Dec 7;186(25):5472-5485.e9.

doi: 10.1016/j.cell.2023.10.018.

Authors

Affiliations

¹ BIOMICs Research Group, Department of Zoology and Animal Cell Biology, University of the Basque Country UPV/EHU, Vitoria-Gasteiz, Spain; Ikerbasque-Basque Foundation of Science, Bilbao, Spain; Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA; Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, Spain. Electronic address: inigo.olalde@ehu.eus.
² Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, Spain.
³ Institute of Archaeology, Belgrade, Serbia.
⁴ Department of Genetics, Harvard Medical School, Boston, MA, USA.
⁵ Department of Genetics, Harvard Medical School, Boston, MA, USA; Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA.
⁶ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA; Department of Genetics, Harvard Medical School, Boston, MA, USA.
⁷ Department of Genetics, Harvard Medical School, Boston, MA, USA; Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA.
⁸ Archaeological Museum Zadar, Zadar, Croatia.
⁹ Department of Archaeology, Durham University, Durham, UK.
¹⁰ BIOMICs Research Group, Department of Zoology and Animal Cell Biology, University of the Basque Country UPV/EHU, Vitoria-Gasteiz, Spain.
¹¹ Slovak National Museum-Archaeological Museum, Bratislava, Slovak Republic.
¹² Centre for Applied Bioanthropology, Institute for Anthropological Research, Zagreb, Croatia.
¹³ Palisada Ltd., Split, Croatia.
¹⁴ School of History, Classics and Archaeology, University of Edinburgh, Edinburgh, UK.
¹⁵ Archaeological Museum in Zagreb, Zagreb, Croatia.
¹⁶ Faculty of Humanities and Social Sciences, University of Zagreb, Zagreb, Croatia.
¹⁷ Division for Archaeological Heritage, Croatian Conservation Institute, Zagreb, Croatia.
¹⁸ Center of Bone Biology, Faculty of Medicine, University of Belgrade, Belgrade, Serbia.
¹⁹ Museum of Vojvodina, Novi Sad, Serbia.
²⁰ Institute of Archaeology, Zagreb, Croatia.
²¹ Department for Archaeology, Croatian Conservation Institute, Zagreb, Croatia.
²² Archaeology Division, Croatian Academy of Sciences and Arts, Zagreb, Croatia.
²³ Museum of Croatian Archaeological Monuments, Split, Croatia.
²⁴ Department of Anthropology and Middle Eastern Cultures, Mississippi State University, Starkville, MS, USA.
²⁵ Department of Anthropology, University of Nevada, Las Vegas, NV, USA.
²⁶ Kaducej Ltd., Split, Croatia.
²⁷ Independent Researcher, 10000 Zagreb, Croatia.
²⁸ Vinkovci Municipal Museum, Vinkovci, Croatia.
²⁹ Trogir Town Museum, Trogir, Croatia.
³⁰ Department of Anthropology, University of Kansas, Lawrence, KS, USA.
³¹ Department of Anthropology, University of Winnipeg, Winnipeg, MB, Canada.
³² Department of Anthropology, Slovak National Museum-Natural History Museum, Bratislava, Slovak Republic.
³³ Laboratory for Bioarchaeology, Faculty of Philosophy, University of Belgrade, Belgrade, Serbia.
³⁴ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria; Department of Anthropology, Natural History Museum Vienna, Vienna, Austria.
³⁵ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria.
³⁶ Servizio di Bioarcheologia, Museo delle Civiltà, Rome, Italy.
³⁷ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria; Research Centre for Anthropology and Health (CIAS), Department of Life Sciences, University of Coimbra, Coimbra, Portugal.
³⁸ Faculty of Biology, University of Belgrade, Belgrade, Serbia; Serbian Academy of Sciences and Arts, Belgrade, Serbia.
³⁹ Faculty of Biology, University of Belgrade, Belgrade, Serbia.
⁴⁰ Department of Classics and Letters, University of Oklahoma, Norman, OK, USA; Santa Fe Institute, Santa Fe, NM, USA.
⁴¹ Department of History, Harvard University, Cambridge, MA, USA; Max Planck-Harvard Research Center for the Archaeoscience of the Ancient Mediterranean, Harvard University, Cambridge, MA, USA.
⁴² Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria; Human Evolution and Archaeological Sciences, University of Vienna, Vienna, Austria.
⁴³ Faculty of Biology, University of Belgrade, Belgrade, Serbia; Department of Biology, University of Western Ontario, London, ON, Canada; Department of Agriculture and Food, Universidad de La Rioja, Logroño, Spain.
⁴⁴ Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, Spain; Museu de Ciències Naturals de Barcelona, Barcelona, Spain. Electronic address: carles.lalueza.fox@gmail.com.
⁴⁵ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA; Department of Genetics, Harvard Medical School, Boston, MA, USA; Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA. Electronic address: reich@genetics.med.harvard.edu.

PMID: 38065079
PMCID: PMC10752003
DOI: 10.1016/j.cell.2023.10.018

Abstract

The rise and fall of the Roman Empire was a socio-political process with enormous ramifications for human history. The Middle Danube was a crucial frontier and a crossroads for population and cultural movement. Here, we present genome-wide data from 136 Balkan individuals dated to the 1^st millennium CE. Despite extensive militarization and cultural influence, we find little ancestry contribution from peoples of Italic descent. However, we trace a large-scale influx of people of Anatolian ancestry during the Imperial period. Between ∼250 and 550 CE, we detect migrants with ancestry from Central/Northern Europe and the Steppe, confirming that "barbarian" migrations were propelled by ethnically diverse confederations. Following the end of Roman control, we detect the large-scale arrival of individuals who were genetically similar to modern Eastern European Slavic-speaking populations, who contributed 30%-60% of the ancestry of Balkan people, representing one of the largest permanent demographic changes anywhere in Europe during the Migration Period.

Keywords: Balkan Peninsula; Great Migration Period; Slavic migrations; ancient DNA; archaeogenetics; cosmopolitanism; demographic changes; population dynamics; the Roman Empire.

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Conflict of interest statement

Declaration of interests The authors declare no competing interests.

Figures

**Figure 1.. Overview of ancient Balkan individuals analyzed in this study.**
(A) Chronological distribution. Individuals with newly reported data are represented by symbols with a black outline. (B) Geographical location of archaeological sites. (C) PCA of the West-Eurasian genetic variability showing present-day individuals as grey circles (except present-day Balkan populations that are displayed with open colored triangles), and relevant ancient populations as colored polygons (Balkan Iron Age groups in blue, Southern European Iron Age groups in light blue, ancient Near Easter groups in red and ancient Steppe, Central, Northern and Eastern European groups in green) including all individuals in each population (Data S2, Table 3). Ancient individuals were projected onto the PCs computed on present-day West Eurasians; their shape and color are the same as in panels (A) and (B). This PCA is a zoom-in version of Figure S1. (D) Closer view of the present-day Balkans genetic cline from panel (C). CNE: Central/Northern European; CEE: Central/Eastern European; BA: Bronze Age; IA: Iron Age.

**Figure 2.. A diversity of ancestral origins.**
(A) By-individual estimates of Iron Age Balkan, West Anatolian/Levantine, and African-related ancestry proportions between 0-1500 CE, computed with *qpAdm*. Two pairs of individuals buried in the same sarcophagi at Rit Necropolis (*Viminacium*) are connected through black lines. (B) δ15N and δ13C stable isotope values (Data S2, Table 9) of ancient Balkan individuals between 0-500 CE obtained from tooth roots, plotted alongside published environmental data and humans from related geographic and chronological contexts^,-. Individuals buried at the same necropolis are connected through lines. (C) Oil lamp depicting an eagle found on individual G-103’s (I15499) grave at Pirivoj, *Viminacium*. (D) Sarcophagus of grave 148 at Rit Necropolis, *Viminacium*.

**Figure 3.. Arrival of ancestry related to Eastern European populations after 700 CE.**
(A) PCA computed on present-day Central, Northern and Eastern Europeans. Present-day Balkan individuals and ancient individuals were projected onto the PCs. Ancient Balkan individuals are shown as red and blue diamonds and other relevant ancient populations are shown as colored polygons including all individuals in each population. (B) f₄-statistics assessing differential affinities to Central/Northern- and Eastern European-related groups. *Central/Northern European-related* includes individuals from two Langobard-associated cemeteries in Hungary and Northern Italy displaying Central/Northern European-related ancestry (*CNE_EarlyMedieval*) and Bronze and Iron Age individuals from the Netherlands (Data S2, Table 3). *Eastern European-related* includes *CEE_EarlyMedieval* and Bronze Age individuals from Latvia and Lithuania. Test populations are shown in the y-axis. Error bars represent one standard error.

**Figure 4.. Demographic impact of Migration Period and Early Medieval events.**
(A) Changes in Central/Northern European, Pontic-Kazakh Steppe and Eastern European-related ancestry proportions between 0-1500 CE, computed with *qpAdm*. A mother and her son are connected through a red line. (B) Proportions of Eastern-European-related ancestry (in black) for present-day Balkan and Aegean populations.

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References

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A genetic history of the Balkans from Roman frontier to Slavic migrations

Affiliations

A genetic history of the Balkans from Roman frontier to Slavic migrations

Authors

Affiliations

Abstract

Conflict of interest statement

Figures

References

Publication types

MeSH terms

Grants and funding

LinkOut - more resources

Full Text Sources