Population history of Early Medieval Ukraine

Lehti Saag1, Olga Utevska2, Stanislav Zadnikov2, Irina Shramko2, Mari Järve3,
Christiana Lyn Scheib , Kyriaki Anastasiadou , Monica Kelly , Mia Williams ,
3, 4 5 5 5

Alexandre Gilardet5, Pontus Skoglund5*, Mark G. Thomas1*

1University College London, 2V. N. Karazin Kharkiv National University,
3University of Tartu, 4University of Cambridge, 5The Francis Crick Institute

*These authors contributed equally to this study

Introduction and Aims of the study

Ukraine is an interesting but so far understudied region in
terms of population history. It borders with Poland, Slovakia
and Hungary to the west while Southern Ukraine is part of
the vast Eurasian Steppe. The region has been in the path
of several migrating groups, including Yamnaya Steppe
pastoralists, mixed-origin Scythians, Gothic Chernyakhov
people, Iranian Alans, Golden Horde Mongols, Turkic
Nogais, and Slavic Cossacks.
This study aims to characterize the genetic affinities of
individuals associated with the Alan group of the Early
Medieval Saltovo-Mayaki Culture, as well as of individuals Ukraine_realcoord_EAA2022

associated with preceding and succeeding cultures. Chernyakhov

Alans
Golden Horde
A modern
modern Nogai
Siberia
Siberia
East Asia
East Asia
Figure 1. Map of Ukrainian sites, samples and
West/Central Asia
West/Central
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Taj TajTuk Uzb
Uzb
UzbUzb Uzb
Uzb
Ala Uyg Kyr Kal Kham Oro
Kal Nan
Ulc
Nivh
Ulc
Ulc
Nan
0.24
0.22
Kum um
Oss
eyOss
yTurum Tur
Tur
Oss Taj Uzb Uzb UygHaz
Haz Oro Nivh
Nan
Ulc 0.23 0.24 0.25 0.26 0.27 0.28
Ady TurTajTaj
Taj Tuk
Uzb Uzb Haz Haz
Haz Nog Oro Ulc
Nivh
Nivh
Nan
Nivh
Oss Bal Taj Uyg
Uyg
UygUyg Haz
Haz
Uyg Nivh
Oro
Oro
Nan
Oro
e
Aze
ur r TurTurUzb −0.02 Uyg
Uyg
Uyg
HazUyg
0.00 0.02 Nog Ulc
Nivh
Nivh
Ulc
0.04
zeTur
r Tur Uyg Uyg
Haz Haz Nivh
Turur
Tur Tur Pat Haz
Haz 0.26
e
ze
ur Tur Pat Tuk Tuk
Tur
ur Tur Tur Pat
Pat Pat Bur
Bur Haz Uyg
Uyg
Oro
ur e PatPat
Pat
Pat Bur
BurBur PC1 (4.2%) Ulc
ze e
Aze
zeAze Pat
Pat Bur
Bur
Bur
Aze
ze
a
e Pat
Pat
Pat
Pat Bur
Bur
B Pat Bur
Pat Bur
Bur
1.0

a Blo Pat
Sin Bur
Bur Oro
abze
aara
Ira
aIra
Sin
Sin Bur
Sin
Sin
Sin
Sin
Bur
Bur
Bur 0.23
aaaa Bur Dun
raBloBlo
Blo Bur Dun 0.25
0.8

aaa Blo BraBloSin

Sin
Sin
Sin DunDun
Dun
– f3(Mbuti; UkrAlan2, Z) –>

Bra
Bra
Blo Dun
aaIra Mak
Blo
Bra
Blo
Bra
Blo
Blo Sin
Bra Sin
Sin
Sin Dun
Dun
Dun
Jap
Mak
MakBlo
Bra
Bra
BloBlo
Blo BloSin Ben HanJap
Jap
Han Jap
Jap
Jap
Bra Bra
0.6

Bra
Mak
Mak
Bra
Mak Blo Ben Han
Dun Han Jap
Jap
JapJap
Jap
Han
MakBlo
BloBlo Ben Jap
J
Jap
Jap
Han ap
Jap
Mak
Mak
Mak
Mak
Bra HanJap
Han
HanJap
Jap
Jap
Mak
Mak
MakBra
Mak Ben
Ben
Ben Han
Han
Han
Han
Mak Ben Han
Han
Han
Han
0.4

Mak Han
Han
Mak
Han
Han
Han
HanHan
Han
Han
Han
Han
Han 0.24 – f3(Mbuti; UkrAlan2, Z) –>
Han
Han
Han
Han 0.22
Han
0.2

0.23 0.24 0.25 0.26 0.27 0.28

0.0

UkrIA_Chernyakhov_0
UkrIA_Chernyakhov_1
UkrIA_Chernyakhov_2
UkrIA_Chernyakhov_3
Sintashta

Poland_BellBeaker
Russia_Siberia_Lena_EBA

China_AmurRiver_N
Russia_Ekven_IA

WHG
China_YR_LN

CHG

Anatolia_N

EHG
SHG

Central_MN

Central_CWC

Ukraine_Crimea_Pantikapaion
Mongolia_North_N

Levant_N

Estonia_CWC

Estonia_EIA

Ukraine_Trypillia

Ukraine_BA
Iran_N

LBK_EN

Estonia_LBA

HungarianScythian

Estonia_MA

Ukraine_GlobularAmphora

Ukraine_LateSrubnaya
Russia_MN_Boisman

Mongolia_Arkhangai_Xiongnu

Hungary_Langobard

Ukraine_Scythian
Russia_Viking

Ukraine_Viking

UkrEMA_Alans_Saltov_1
UkrEMA_Alans_Saltov_2
UkrEMA_Alans_Saltov_3
Mongolia_EIA_SlabGrave_1

Poland_GlobularAmphora

Ukraine_Mesolithic
Ukraine_Neolithic

Ukraine_Eneolithic
Ukraine_Yamnaya

UkrMA_GoldenHorde
Yamnaya_Kalmykia
Yamnaya_Samara

TianShanHun

Fatyanovo

Sweden_Viking

Ukraine_Cimmerians

UkrMA_Nogai_1
UkrMA_Nogai_2
UkrMA_Nogai_3
Serbia_Mokrin_EBA_Maros

BolshoyOleniOstrov
Czech_BellBeaker

UkrMA_GoldenHorde_Slav-Tat
Scythian_East

Ukraine_Medieval

Ukrainian_proj
Germany_EarlyMedieval
Poland_Medieval
Italy_Medieval

– f3(Mbuti; UkrAlan1, Z) –>

0.23
0.21 0.22 0.23 0.24 0.25 0.26 0.27

Figure 3. f3 statistics results comparing genetic

subgroups of Alan-associated individuals.
0.00 0.02 0.04
Results and Conclusion
1.0

Here we present novel genome-wide shotgun sequencing

PC1 (4.2%)
data from 30 individuals – 8 associated with the Alan group
0.8

of the Early Medieval Saltovo-Mayaki Culture, 11

0.6

associated with the preceding Chernyakhov Culture and 4

0.4

and 7, respectively, from the succeeding Golden Horde and

0.2

Nogai periods. The genomes have been sequenced to an

Poland_Medieval0.0

average coverage of 0.38x (0.02–0.95x).

UkrIA_Chernyakhov_0
UkrIA_Chernyakhov_1
UkrIA_Chernyakhov_2
UkrIA_Chernyakhov_3

UkrIA_Chernyakhov_0
UkrIA_Chernyakhov_1
UkrIA_Chernyakhov_2
UkrIA_Chernyakhov_3
Sintashta

Poland_BellBeaker
Russia_Siberia_Lena_EBA

China_AmurRiver_N
Russia_Ekven_IA

China_YR_LN

CHG

Anatolia_N

Ukraine_Crimea_Pantikapaion

EHG
SHG
WHG

Central_MN

Central_CWC

Ukraine_Crimea_Pantikapaion
Mongolia_North_N

Ukraine_BA

Ukraine_BA
Estonia_MA

Ukraine_GlobularAmphora
Ukraine_Trypillia

Ukraine_LateSrubnaya

Levant_N

Estonia_CWC

Estonia_LBA

Estonia_EIA

Ukraine_LateSrubnaya
Hungary_Langobard

Ukraine_Scythian
Russia_Viking

Iran_N

Ukraine_Viking

UkrEMA_Alans_Saltov_1
UkrEMA_Alans_Saltov_2
UkrEMA_Alans_Saltov_3
LBK_EN

HungarianScythian

Estonia_MA

Ukraine_GlobularAmphora
Ukraine_Trypillia

Ukraine_Scythian

Ukraine_Viking

UkrEMA_Alans_Saltov_1
UkrEMA_Alans_Saltov_2
UkrEMA_Alans_Saltov_3
Russia_MN_Boisman
Ukraine_Mesolithic
Ukraine_Neolithic

Ukraine_Eneolithic
Ukraine_Yamnaya

Mongolia_Arkhangai_Xiongnu

UkrMA_GoldenHorde

Hungary_Langobard

Russia_Viking
Poland_GlobularAmphora
UkrMA_Nogai_1
UkrMA_Nogai_2
UkrMA_Nogai_3

Ukraine_Mesolithic
Ukraine_Neolithic

Ukraine_Eneolithic
Ukraine_Yamnaya

UkrMA_GoldenHorde
Sweden_Viking

Mongolia_EIA_SlabGrave_1

Ukraine_Cimmerians

Yamnaya_Kalmykia

Ukraine_Cimmerians

UkrMA_Nogai_1
UkrMA_Nogai_2
UkrMA_Nogai_3
Yamnaya_Samara

TianShanHun

Fatyanovo

Sweden_Viking
Serbia_Mokrin_EBA_Maros

BolshoyOleniOstrov
UkrMA_GoldenHorde_Slav-Tat

Czech_BellBeaker

UkrMA_GoldenHorde_Slav-Tat
Scythian_East
Ukraine_Medieval

Ukrainian_proj

Ukraine_Medieval

Ukrainian_proj
Germany_EarlyMedieval

Germany_EarlyMedieval
Poland_Medieval
Italy_Medieval

Italy_Medieval

The first results show variability in genetic affinities among

individuals with the same cultural association (especially
within Alan-associated individuals), as well as between
different groups and time periods. Genetic influences from
the Near East, the Pontic-Caspian steppe, and East Asia
Figure 2. PCA (A) and ADMIXTURE analysis (B) results. can be inferred.

Contacts Acknowledgements
l.saag@ucl.ac.uk pontus.skoglund@crick.ac.uk m.thomas@ucl.ac.uk This project has received funding from the European Union’s
@LehtiSaag @pontus_skoglund @mt_genes Horizon 2020 research and innovation programme under the
Marie Skłodowska-Curie grant agreement No 101029303.