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Our Avar period samples also form a characteristic PCA “Avar-cline” on Fig. 2,
extending from Europe to Asia. PC50 clustering identified a single genetic cluster at the Asian
extreme of the cline with 12 samples, derived from 8 different cemeteries, which we termed
Avar_Asia_Core (Fig. 2, Supplementary Table 3). 10/12 samples of Avar_Asia_Core were
assigned to the early Avar period, 4 of them belonging to the elite, and 9/12 were males.
Avar_Asia_Core clusters together with Shamanka_Eneolithic and
Lokomotiv_Eneolithic19 samples from the Baikal region, as well as with Mongolia_N_East,
Mongolia_N_North15
, Fofonovo_EN, Ulaanzuukh_SlabGrave and Xiongnu14 from Mongolia
(Supplementary Table 3). This result is recapitulated in ADMIXTURE (Fig. 2b), which also
shows that Nganasan and Han components predominate in Avar_Asia_Core with traces of
Anat_N and ANE, while Iranian and WHG ingredients are entirely missing. It follows, that
Avar_Asia_Core was derived from East Asia, most likely from present day Mongolia.
We performed two-dimensional f4-statistics to detect minor genetic differences within
the Avar_Asia_Core group. Avar_Asia_Core individuals could be separated along a Bactria-
Margiana Archaeological Complex (BMAC)- Steppe Middle-Late Bronze Age
(Steppe_MLBA) cline (Extended Data Fig 3), with 3 individuals bearing negligible
proportion of these ancestries. The Steppe_MLBA-ANE f4-statistics gave similar results. As
the 3 individuals with the smallest Iranian, Steppe and ANE ancestries also visibly separated
on PCA, we set apart these under the name of Avar_Asia_Core1, while the other 9 samples
were regrouped as Avar_Asia_Core2 (Fig. 2).
According to outgroup f3-statistics both Avar_Asia_Core groups had highest shared
drift with genomes having predominantly Ancient North-East Asian (ANA) ancestry
(Extended Data Fig. 2), like earlyXiongnu_rest, Ulaanzuukh, and Slab Grave14
. It is notable
that from the populations with top 50 f3 values, 41 are shared with Hun_Asia_Core, moreover
Avar_Asia_Core1 is the 16th in the top list of VZ-12673 and 35th in that of MSG-1, signifying
common deep ancestry of European Huns and Avars.
According to distal qpAdm models Avar_Asia_Core formed a clade with the
Fofonovo_EN and centralMongolia_preBA genomes (Supplementary Table 6a), both of
which had been modelled from 83%–87% ANA and 12%–17% ANE14
. All data consistently
show that Avar_Asia_Core preserved very ancient Mongolian pre-Bronze Age genomes, with
~90% ANA ancestry.
Most proximodistal qpAdm models (defined in Methods) retained distal sources, as
Avar_Asia_Core1 was modelled from 95% UstBelaya_N20 plus 5% Steppe Iron Age
(Steppe_IA) and Avar_Asia_Core2 from 80-92% UstBelaya_N plus 8-20% Steppe_IA
(Supplementary Table 6b). The exceptional proximal model for Avar_Asia_Core1 indicated
58% Yana_Medieval20 plus 42% Ulaanzukh, while for Avar_Asia_Core2 69%
Xianbei_Hun_Berel13 plus 31% Kazakhstan_Nomad_Hun_Sarmatian12 ancestries. The latter
model also points to shared ancestries between Huns and Avars.
From the 76 samples in the Avar-cline, 26 could be modelled as a simple 2-way
admixture of Avar_Asia_Core and EU_Core (Supplementary Table 6c) indicating that these
were admixed descendants of locals and immigrants, while further 9 samples required
additional Hun and/or Iranian related sources. In the remaining 40 models Hun_Asia_Core
and/or Xiongnu sources replaced Avar_Asia_Core (Supplementary Table 6d, summarized in
Supplementary Table 1b). Scythian-related sources with significant Iranian ancestries, like
Alan, Tian Shan Hun, Tian Shan Saka12
, or Anapa (this study), were ubiquitous in the Avar-
cline, but given their low proportion, qpAdm was unable to identify the exact source.
Xiongnu/Hun-related ancestries were more common in certain cemeteries, for example
it was detected in most samples from Hortobágy-Árkus (ARK), Szegvár-Oromdűlő (SZOD),
Makó-Mikócsa-halom (MM) and Szarvas-Grexa (SZRV). Y-chromosomal data seem to
corroborate this conclusion, as 8/10 males from ARK carried Y-Hg Q, while 2/10 R1a-Z94,
3/3 males from SZRV carried R1a-Z94 and 2/2 males from MM carried Hg Q
(Supplementary Table 1a).