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higher resolution using earlier genomes. Several other individuals from 
these Longobard burials (Longobard_earlyMED(II)) show no detectable 
ancestry from northern Europe and, instead, are more closely related to 
Iron Age groups in continental central Europe, putatively representing 
descendants of local people buried in a Longobard style. Our results are 
consistent with attestations that the Longobards originated in the areas 
of present-day northern Germany or Denmark, but that by the sixth 
century ce they incorporated multiple different cultural identities, and 
mixed ancestries. Present-day populations of Hungary do not appear 
to derive detectable ancestry from early medieval individuals from 
Longobard contexts, and are instead more similar to Scythian-related 
ancestry sources (Extended Data Fig. 6), consistent with the later impact 
of Avars, Magyars and other eastern groups58.
In southern Germany, the genetic ancestry of individuals from 
early medieval Bavaria probably associated with the historical 
Germanic-language-speaking Baiuvarii59 cannot be modelled as deriv-
ing ancestry solely from earlier groups in Iron Age central Germany 
(P ≪ 1 × 10−36). The Baiuvarii probably appeared in the region in the 
fifth century ce59, but their origins remain unresolved. Our current 
best model indicates a mixture with ancestry derived from EIA Pen-
insular Scandinavia and central Europe, suggesting an expansion of 
Scandinavian-related ancestry producing a regional ancestry shift 
(Figs. 2c and 3c).
In Italy, southward expansions of northern and central European ances-
tries appear by the Late Antiquity (approximately fourth century ce),  
where a clear diversification of ancestry can be observed compared 
with preceding time periods (Fig. 3d). However, no individuals with 
near 100% Scandinavian ancestry can be observed in the sampling 
data available so far.
In Britain, the ancestries of Iron Age and Roman individuals form a 
tight cluster in our MDS analysis (Fig. 3e), shifted relative to available 
preceding Bronze Age individuals from Ireland and Orkney, and adja-
cent to, but distinct from, available individuals in Iron Age and Roman 
central Europe. However, two first- to second-century ce burials from a 
Roman military fortress site in Austria (Klosterneuburg)5 carry ancestry 
that is currently indistinguishable from Iron Age or Roman popula-
tions of Britain, to the exclusion of other groups (qpWave cladality  
P = 0.11). One option is that they had ancestry from Britain; alternatively, 
currently unsampled populations from western continental Europe 
carried ancestries similar to Iron Age southern Britain.
Twigstats substantially improves models of admixture between 
ancestries from Iron Age Britain and northern Europe in early medi-
eval England9, halving standard errors from 9% with SNPs to 4% when 
using time stratification (point estimates 80% and 79% Iron Age 
Britain-related ancestry, respectively). We used this improved reso-
lution to demonstrate that an earlier Roman individual (6DT3) dating 
to approximately second to fourth century ce from the purported 
gladiator or military cemetery at Driffield T errace in York (Roman 
Eboracum), England60, who was previously identified as an ancestry 
outlier61,62, specifically carried approximately 25% EIA Scandinavian 
Peninsula-related ancestry (Fig. 2c). This documents that people with 
Scandinavian-related ancestry already were in Britain before the fifth 
century ce, after which there was a substantial influx associated with 
Anglo-Saxon migrations9. Although it is uncertain whether this indi-
vidual was a gladiator or soldier, individuals and groups from northern 
Europe are indeed recorded in Roman sources both as soldiers and as 
enslaved gladiators63,64.
Across Europe, we see regional differences in the southeastern and 
southwestern expansions of Scandinavian-related ancestries. Early