Shift in the genetic landscape of the western Eurasian Steppe not due to Scythian

dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths)

Mari Järve1*, Christiana Lyn Scheib1, Lehti Saag1, Kristiina Tambets1, Lauri Saag1, Luca Pagani1, Ajai K. Pathak1, J. Rodrigo Flores1, Francesco
Montinaro1, Liivi Varul2, Stanislav Zadnikov3, Oleg Petrauskas4, Maryana Avramenko4, Boris Magomedov4, Serghii Didenko5, Gennadi Toshev6, Igor
Bruyako7, Denys Grechko4, Vitali Okatenko8, Kirill Gorbenko9, Aleksandr Smyrnov9, Anatolii Heiko10, Roman Reida4, Serheii Sapiehin11, Sergey Sirotin12,
Aleksandr Tairov13, Arman Beisenov14, Maksim Starodubtsev15, Vitali Vasilev16, Alexei Nechvaloda17, Biyaslan Atabiev18, Sergey Litvinov17, Murat
Dzhaubermezov19, Olga Utevska3, Irina Shramko3, Elza Khusnutdinova17,19, Nikita Savelev17, Aivar Kriiska1, Toomas Kivisild1,20, Richard Villems1
1. University of Tartu, Tartu, Estonia; 2. Tallinn University, Tallinn, Estonia; 3. V. N. Karazin Kharkiv National University, Kharkiv, Ukraine; 4. National Academy of Sciences of Ukraine, Kyiv, Ukraine; 5. National Museum of History
of Ukraine, Kyiv, Ukraine; 6. Zaporizhia National University, Zaporizhia, Ukraine; 7. Odessa Archaeological Museum, Odessa, Ukraine; 8. National Academy of Sciences of Ukraine, Kharkiv, Ukraine; 9. Mykolaiv National University,
Mykolaiv, Ukraine; 10. National Museum of Ukrainian Pottery, Opishne, Ukraine; 11. Poltava Regional Makarenko Scientific Lyceum, Kovalivka, Ukraine; 12. Russian Academy of Sciences, Moscow, Russia; 13. South Ural State
University, Chelyabinsk, Russia; 14. A. H. Margulan Institute of Archaeology, Almaty, Kazakhstan; 15. Sterlitamak Krai Museum, Sterlitamak, Russia; 16. LoCom Medien Akademie Europäisches Bildungsinstitut, Bonn, Germany; 17.
Ufa Federal Research Centre of the Russian Academy of Sciences, Ufa, Russia; 18. Institute of the Caucasus Archaeology, Nalchick, Russia; 19. Bashkir State University, Ufa, Russia; 20. University of Cambridge, Cambridge, UK
* Corresponding author: mari.jarve@ut.ee

THE SCYTHIANS
The Early Iron Age nomadic Scythians have been shown to have been a
confederation of tribes of different origins based on ancient DNA b Sumbar_LBA
Tepe_Hissar_C
Shahr_I_Sokhta_BA3
Shahr_I_Sokhta_BA2
Shahr_I_Sokhta_BA1

evidence1,2. It is still unclear how much of the Scythian dominance in the Sappali_Tepe_BA
Gonur2_BA
Gonur1_BA
Parkhai_LBA

Eurasian Steppes was due to movements of people and how much reflected Parkhai_MBA
Parkhai_EBA
Sarazm_EN
Tepe_Anau_EN

cultural diffusion and elite dominance. Parkhai_EN

Geoksiur_EN
Ganj_Dareh_N
Bustan_BA

We present new whole-genome results of 31 ancient samples (Figure 1), Dzharkutan1_BA

Darra_i_kur_MBA
Dzharkutan2_BA
Hajji_Firuz_C

dividing them into 6 groups to facilitate comparisons: Hajji_Firuz_BA

Kanai_MBA
NLithuania
Lithuania_BA
Latvia_BA

preScy_Ukr Ukrainian pre-Scythians 6 samples Lithuania_CWC

Latvia_CWC
Estonia_CWC
Latvia_CCC_EHG

Scy_Ukr Ukrainian Scythians 9 samples Estonia_CCC

Latvia_CCC_WHG
Lithuania_HG
Latvia_HG

Chern Chernyakhov (Ukraine) 3 samples Estonia_HG

Ukraine_Eneolithic
Hungarian_Med
Sweden_LNBA

Sar_Cau Caucasus Sarmatian 1 sample Northern_LNBA

Hungary_BA
HallstattBylany
Central_LNBA

ScySar_SU S Urals Scythians/Sarmatians 8 samples Bell_Beaker

Trypillia
Sweden_TRB
Remedello

Scy_Kaz Kazakhstan Scythians 4 samples LBK_EN

Iberia_MN
Iberia_EN
Iberia_Chl
Iberia_BA
Hungary_EN
Globular_Amphora
Central_MN
Saa Cardial_EN
WHG
Iron_Gates_HG
Bichon
SHG
Kha
Kar Romania_HG
Lat Vep Ukraine_Neolithic
Lit Fin SelKet
Ukraine_Mesolithic

Est Nen SidelkinoEHG_ML

EHG
Kom Ngn
NRu Udm Mar
Nomad_His
Karakhanid
Pol CRu Kipchak
Blr Mrd Karluk
SRu Chv Kimak
ChkKor
Swe
Ukr Figure 1. Map of sample locations + timeline. Nomad_Med
Nomad_Hun-Sarmatian
Nomad_HP
Turk
Slk Ttr Evn TianShanHun
Orc
DIFFERENT SCYTHIAN GROUPS
XiongNu_WE
Ger Dol XiongNu
Sln
Hun Bsh Ykg GoldenHordeAsian
Shr GoldenHordeEuro
Evk Kazakh_His
Bos
FrBFre
Srb
Khk
Yak
Modern population medians We can confirm the confederate nature SaltovoMayaki
Kangju
Wusun
Central Saka Scy_Kaz

Alt
Tuv
Hungarian Scythians of Scythian tribes: the Scy_Ukr, Scy_Kaz
Scy_Kaz
PC2 (0.65%)

Mac Scy_Kaz

samples of this study

Rom
Bul ScySar_SU
Hun-Sarmatian Nomads

NIt
Gag Kaz
Kal
Bur
Iron Age Nomads
ScySar_SU and Scy_Kaz groups are ScySar_SU
ScySar_SU
ScySar_SU
Kyr Mon ScySar_SU

Tus Nog Uzb

Orq Sarmatians
clearly distinguishable in PCA (Figure ScySar_SU
ScySar_SU
ScySar_SU
Grk Tagar Sar_Cau

Sar
Tjk Uyg
Dau
Hzh
Tian Shan Saka 2), most likely due to their different Chern
Chern
Chern
Lzg Psh Iron Age Scythian
Che Scy_Ukr

Kum
Blk
Ady NOs MglXib Southern Urals Sarmatians
geographic origins. Expectedly, there is Scy_Ukr
Scy_Ukr
Scy_Ukr

a decreasing gradient of eastern (‘Han’ /

Scy_Ukr
Trm Tu Eastern Scythians Scy_Ukr
Scy_Ukr
Tur Jap preScy_Ukr Scy_Ukr

‘Nganasan’) ancestry from east to west,

Scy_Ukr
Abh Aze Han Scy_Ukr
preScy_Ukr
preScy_Ukr
Krd preScy_Ukr
CypGrg Irn Chern
Arm
Sar_Cau
with Scy_Kaz samples displaying the preScy_Ukr
preScy_Ukr
preScy_Ukr
Scythian_East
Leb
Syr ScySar_SU
Scy_Kaz
highest and Scy_Ukr samples the lowest Scythian_East
Scythian_East
Scythian_East
Scythian_East

Jor proportions (Figure 3). Scythian_East

CentralSaka
CentralSaka
Pal CentralSaka
CentralSaka
CentralSaka

PC1 (4.37%) Figure 3. ADMIXTURE results of CentralSaka

NoDateTianShanSaka
TianShanSaka

a) modern; b) ancient samples. TianShanSaka

TianShanSaka
TianShanSaka

Figure 2. PCA of novel and published ancient samples from Scythian/Sarmatian and related TianShanSaka
TianShanSaka

groups on the background of modern samples presented as population medians. a Chukchis

Koryaks
Evens
Evenkis
TianShanSaka
TianShanSaka
TianShanSaka
TianShanSaka
Yakuts Tagar

∆ - ref. 1, ○ - ref. 2, □ - ref. 3, ◊ - this study. Yukaghirs

Shors
Selkups
Tagar
Tagar
Tagar
Nganasans Tagar
Kets Tagar
Dolgans Tagar

GENETIC SHIFT NOT WITH THE WESTERN SCYTHIANS, BUT AFTER THEM
Tuvinians Tagar
Buryats Nomad_IA
Japanese Nomad_IA
Mongolians Nomad_IA
Nomad_IA
Mongola

In the Ponto-Caspian region, we can set the Scythians in a temporal context by comparing the Oroqens
Hezhen
Daur
Scythian_IA
Sarmatian_SU
Sarmatian_SU
Sarmatian
Xibo

Scy_Ukr samples to preScy_Ukr samples predating and Chern samples postdating them. Population Tu
Han
Sakilli
Sarmatian
Sarmatian
Sarmatian
Sarmatian
Paniya

modelling results (Figure 4), confirmed by f4 test, reveal no difference in the eastern (Nganasan) Malayan
North_Kannadi
Sindhi
Sarmatian
Sarmatian
Sarmatian
Sarmatian

affinity of preScy_Ukr and Scy_Ukr, while Scy_Ukr share significantly more alleles with Nganasan Makrani
Brahui
Balochi
Burusho
HungarianScythian
HungarianScythian
HungarianScythian
HungarianScythian

than Chern do. On the other hand, Chern display the highest proportion of early farmer-related Pashtun
Pathan
Hazara
Khakases
HungarianScythian

Turkmenistan_IA
Poprad

Zevakinskiy_LBA

(Anatolian Neolithic) ancestry among the sample groups of this study (Figure 4). Altaians
Uygurs
Kazakhs
Zevakinskiy_MLBA
Zevakinskiy_BA
Srubnaya
Kyrgyzians Sintashta_MLBA

Thus we detect no significant contribution of the Scythians to the Ponto-Caspian gene pool where Uzbeks
Tajiks
Turkmens Preobrazhenka_MLBA
Sintashta
Petrovka

Kalmyks Alpamsa_MLBA_Alakul

no large-scale changes appear to occur from the Yamnaya through the Scythian period (~4,800– Nogais
Kumyks
Lezgins
Molaly_MLBA
Taldysay_MLBA2
Taldysay_MLBA1
Chechens Solyanka_MLBA

2,100 cal BP). However, the transition from the Scythian to the Chernyakhov culture (~2,100–1,700 North_Ossetians
Adygei
Balkars
Kyzlbulak_MLBA2
Kyzlbulak_MLBA1
Oy_Dzhaylau_MLBA
Abkhasians Satan_MLBA_Alakul

cal BP) does mark a shift in the Ponto-Caspian genetic landscape. Georgians
Armenians
Tatars
Ak_Moustafa_MLBA1
Maitan_MLBA_Alakul
Lisakovskiy_MLBA_Alakul
Bashkirs Krasnoyarsk_MLBA

Our results agree well with the Ostro- Chuvash

Hantis
Nenets
Kazakh_Mys_MLBA
Kashkarchi_BA
Karagash_MLBA
Udmurts Kairan_MLBA

gothic origins of the Chernyakhov culture Komis

Mordovians
Maris
Dashti_Kozy_BA
Dali_MLBA
Andronovo
Aktogai_MLBA

and support the hypothesis that Scythian Vepsas

Saami
Swedes
Karelian
Yamnaya_Ukraine
Yamnaya_Samara
Yamnaya_Kalmykia
Yamnaya_Bulgaria

dominance was cultural rather than Finns

Estonians
Latvians
Lithuanians
Yamnaya
Russia_EBA
Potapovka
Poltavka

achieved through population replacement. Russians_North

Russians_Central
Russians_South
Okunevo_EMBA
CentralSteppe_EMBA
Glazkovo
Belarusians
Kurma_EBA
Ukrainians
Shamanka_EBA
Poles
UstIda_LN

Figure 4. ChromoPainter/NNLS mixture fit result of the

Slovaks
Lokomotiv_EN
Gagauzes
Shamanka_EN
Romanians
Afanasievo
Hungarians

sample groups of this study as a combination of Eastern

Namazga_CA
Bulgarians Dali_EBA
Slovenians Botai
Serbians Samara_Eneolithic

Hunter-Gatherer (EHG), Anatolian Neolithic (AN), Bosnians

Macedonians
Greeks
West_Siberia_N
Kostenki
Mal'ta
Cypriots

Iranian Neolithic (IN), and Nganasan.

Ust-Ishim
Tuscans Levant_BA
North_Italians Anatolia_ChL
Sardinians Levant_N
French_Basques Anatolia_N
Germans Natufian

REFERENCES
French LchashenMetsamor
Orcadians Alan
Azeris Armenia_MLBA

1. Unterländer, M. et al. Ancestry and demography and descendants of Iron Age nomads of the Eurasian Steppe. Nat Comm 8, Article 14615 (2017). Iranians
Kurds
Armenia_EBA
Iran_ChL
Turks Armenia_ChL
2. Damgaard, P. de B. et al. 137 ancient human genomes from across the Eurasian steppes. Nature 557, 369–374 (2018). Lebanese
Syrians
Iran_LN
Iran_N
Jordanians Iran_HotuIIIb
3. Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419–424 (2016). Palestinians CHG
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