{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T23:00:44Z","timestamp":1784847644341,"version":"3.55.0"},"reference-count":50,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T00:00:00Z","timestamp":1784851200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Genet."],"abstract":"<jats:p>\n                    We report genome-wide ancient DNA from 37 individuals retained after conservative archaeological and genomic reassessment of 53 screened samples from two Gothic-period mortuary contexts in present-day Bulgaria: the Aquae Calidae necropolis in Thrace (n = 20; c. 320 CE\u2013375 CE) and the Aul of Khan Omurtag (AKO) in Moesia Secunda (n = 17; c. 350 CE\u2013489 CE). The filtering process excluded individuals with uncertain or probably later-medieval horizon assignment, including a C5-area component linked by READv2 kinship to suspected later-horizon burials. We evaluate three non-exclusive models for Balkan Gothic formation, namely, migration-continuity from northern and Pontic Gothic horizons, a Wenskus\u2013Wolfram\u2013Pohl tradition-bearing cultural\u2013political formation model across changing demographic substrates, and the Roman frontier formation. The two assemblages share Gothic-associated material culture and Christian east\u2013west burial orientation, but they are not a single genome-wide population. Proximal qpAdm models distinguish a southern Anatolian\/Marmara-related and northern\/Pontic structure at Aquae Calidae from a simpler Chernyakhov-related and Balkan Late Antique structure at AKO. f\n                    <jats:sub>4<\/jats:sub>\n                    statistics further show that the Anatolian-related component at Aquae Calidae cannot be explained solely as a Chernyakhov-carried signal. DATES places north\u2013south admixture at approximately 12.44 \u00b1 2.36 generations before burial (Z = 5.26), consistent with a pre-burial ancestry mixture rather than a single intact Gothic migration. Kinship is confined within sites. Together, the data reject a single biological Gothic population moving unchanged into the Balkans and instead support a formation process combining migration, multiethnic coalition, and frontier incorporation.\n                  <\/jats:p>","DOI":"10.3389\/fgene.2026.1864752","type":"journal-article","created":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T22:10:22Z","timestamp":1784844622000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Paleogenomic evidence for genetic heterogeneity and prior admixture in Gothic-associated communities of late antique Bulgaria"],"prefix":"10.3389","volume":"17","author":[{"given":"Svetoslav","family":"Stamov","sequence":"first","affiliation":[{"name":"Laboratory of Interdisciplinary Research and Paleogenomic Knowledge, National Museum of History","place":["Sofia, Bulgaria"]},{"name":"UNIBIT","place":["Sofia, 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Adv."},{"key":"B40","doi-asserted-by":"crossref","first-page":"300","DOI":"10.17433\/978-3-17-025169-4","volume-title":"Rom Und Die Barbaren: V\u00f6lker Im Alpen-Und Donauraum","author":"Steinacher","year":"2017"},{"key":"B41","doi-asserted-by":"publisher","first-page":"6737","DOI":"10.1038\/s41598-019-43183-w","article-title":"Goth migration induced changes in the matrilineal genetic structure of the central-east European population","volume":"9","author":"Stolarek","year":"2019","journal-title":"Sci. Rep."},{"key":"B42","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1186\/s13059-023-03013-9","article-title":"Genetic history of east-central Europe in the first millennium CE","volume":"24","author":"Stolarek","year":"2023","journal-title":"Genome Biol."},{"key":"B43","doi-asserted-by":"crossref","DOI":"10.5040\/9781472541000","volume-title":"The Visigoths in the Time of Ulfila","author":"Thompson","year":"2008"},{"key":"B44","doi-asserted-by":"publisher","first-page":"3494","DOI":"10.1073\/pnas.1719880115","article-title":"Population genomic analysis of elongated skulls reveals extensive female-biased immigration in early medieval Bavaria","volume":"115","author":"Veeramah","year":"2018","journal-title":"PNAS"},{"key":"B45","doi-asserted-by":"publisher","first-page":"W58","DOI":"10.1093\/nar\/gkw233","article-title":"HaploGrep 2: mitochondrial haplogroup classification in the era of high-throughput sequencing","volume":"44","author":"Weissensteiner","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"B46","doi-asserted-by":"crossref","DOI":"10.7788\/9783412300067","volume-title":"Stammesbildung Und Verfassung: Das Werden Der Fr\u00fchmittelalterlichen Gentes","author":"Wenskus","year":"1961"},{"key":"B47","doi-asserted-by":"publisher","first-page":"iyae110","DOI":"10.1093\/genetics\/iyae110","article-title":"Testing times: disentangling admixture histories in recent and complex demographies using ancient DNA","volume":"228","author":"Williams","year":"2024","journal-title":"Genetics"},{"key":"B48","volume-title":"History of the Goths","author":"Wolfram","year":"1988"},{"key":"B49","article-title":"Typen der Ethnogenese am Beispiel der Bayern","volume":"1","author":"Wolfram","year":"1990","journal-title":"Typen Ethnogenese unter Bes. 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