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A Dynamic 6,000-Year Genetic History of Eurasia’s Eastern Steppe

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A Dynamic 6,000-Year Genetic History of Eurasia’s Eastern Steppe



The Eastern Eurasian Steppe was home to historic empires of nomadic pastoralists, including the Xiongnu and the Mongols. However, little is known about the region’s population history. Here, we reveal its dynamic genetic history by analyzing new genome-wide data for 214 ancient individuals spanning 6,000 years. We identify a pastoralist expansion into Mongolia ca. 3000 BCE, and by the Late Bronze Age, Mongolian populations were biogeographically structured into three distinct groups, all practicing dairy pastoralism regardless of ancestry. The Xiongnu emerged from the mixing of these populations and those from surrounding regions. By comparison, the Mongols exhibit much higher eastern Eurasian ancestry, resembling present-day Mongolic-speaking populations. Our results illuminate the complex interplay between genetic, sociopolitical, and cultural changes on the Eastern Steppe.

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Genetic group ID Time Period Affiliated cultures Number of Samples Sample IDs
eastMongolia_preBA pre-Bronze Age Unknown 1 SOU001
centralMongolia_preBA pre-Bronze Age Unknown 1 ERM001
Fofonovo_EN pre-Bronze Age Fofonovo  4 FNO001; FNO003; FNO006; FNO007
Afanasievo_Mongolia Early Bronze Age Afanasievo 2 SHT001; SHT002
Afanasievo_KUR001 Early Bronze Age Afanasievo 1 KUR001
Chemurchek Early Bronze Age Chemurchek 2 YAG001; IAG001
Chemurchek_KUM001 Early Bronze Age Chemurchek 1 KUM001
Altai_MLBA Middle Late Bronze Age Mönkhkairkhan, DSKC, Baitag 7 SBG001; BER002; ULI003; ULI001; BIL001; KHI001; ARS026*
KHU001 Middle Late Bronze Age Munkhkairkhan 1 KHU001
Ulaanzuukh_SlabGrave Middle Late Bronze Age; Early Iron Age Ulaanzuukh; Slab Grave 16 ULN010;ULN006;ULN009;ULN002;ULN005;BUL002;ULN001;ULN003;ULN007;BUL001;ULN015;DAR001;MIT001;BOR001;SHU001;PTO001
UAA001 Middle Late Bronze Age Munkhkairkhan 1 UAA001
ULZ001 Middle Late Bronze Age Unknown 1 ULZ001
UUS001 Middle Late Bronze Age DSKC 1 UUS001
TSI001 Middle Late Bronze Age Unknown 1 TSI001
ARS017 Middle Late Bronze Age Unknown 1 ARS017
Khovsgol_LBA Middle Late Bronze Age DSKC 17 ARS001,ARS002,ARS003,ARS005,ARS006,ARS007,ARS008,ARS009,ARS011,ARS012,ARS013,ARS014,ARS015,ARS016,ARS018,ARS024,ARS025
Chandman_IA Early Iron Age Uyuk 9 CHN007;CHN016;CHN012;CHN003;CHN008;CHN006;CHN001;CHN015;CHN004
earlyXiongnu_west early Xiongnu Xiongnu 6 SKT010;SKT001;SKT003;SKT009;SKT008;AST001
earlyXiongnu_rest early Xiongnu Xiongnu 6 SKT005;SKT012;SKT006;SKT002;SKT004;JAG001
SKT007 early Xiongnu Xiongnu 1 SKT007
lateXiongnu lateXiongnu Xiongnu 24 SAN001,UGU004,SOL001,DEL001,ULN004,BTO001,CHN010,DOL001,JAA001,KHO006,KHO007,TEV002,TUK003,UGU011,UVG001,TEV003,IMA001,IMA002,IMA003,IMA004,IMA005,IMA006,IMA007,IMA008
lateXiongnu_sarmatian lateXiongnu Xiongnu 13 TMI001,BUR001,NAI001,BUR002,BRL002,BUR003,BUR004,DUU001,HUD001,NAI002,UGU005,UGU006,UGU010
lateXiongnu_han lateXiongnu Xiongnu 8 ATS001,BAM001,BRU001,EME002,SON001,TUH001,TUH002,YUR001
YUR001 lateXiongnu Xiongnu 1 YUR001
TUK001 early Medieval Unknown 1 TUK001
earlyMed_Turk early Medieval Turk 7 ZAA004;TSB001;ULI002;NOM001;ZAA002;UGU001;ZAA007
earlyMed_Uigur early Medieval Uigur 12 OLN011;OLN001.B;OLN003;OLN005;OLN002;OLN008;OLN001.A;OLN010;OLN012;ZAA001;OLN004;OLN009
TUM001 early Medieval probably ancient Chinese 1 TUM001
OLN007 early Medieval Uigur 1 OLN007
lateMed_Khitan late Medieval Khitan 3 ZAA003;ZAA005;ULA001
lateMed_Mongol late Medieval Mongol 61 ARG001;ARG002;ARG003;KHV002;KNN001;CHD001;KRN001;KRN002;TSA001;TSA002;TSA003;MRI001;TSA005;BAY001;TAH002;NRC001;TAV001;TAV006;TAV005;DAR002;DAS001;BRG004;BRG005;KGK001;ZAR002;TSA004;KHN001;KHO001;BAZ001;GTO001;GUN002;BRG001;BRG002;IKU001;SHG002;SHG003;KHL001;UGO001;UGO002;UGU002;UGU003;RAH001;SHG001;ZAY001;KNU001;ZAM002;TSA007;SHR001;GAN002;TSA006;ERD001;TAV011;SHA001;ZAM001;ULN011;CHN014;DEE001;BAU001;BRL001;DUU002;UUS002
SHU002 late Medieval Unknown 1 SHU002


ARG001 M G2a1 R1a1a1b2a2b (R-Z2122)
ARG002 F M10a2 -
ARG003 M C4a2a1 C2b1a1b1 (C-F3830)
ARS001 M D4 Q1a2a1c (Q-L334; Q-L330)
ARS002 F D4j5 -
ARS003 M U5a2d1 N1c1a (N-M178)
ARS004 M F2a Q1a2 (Q-L942; Q-M346)
ARS005 M D4c1 Q1a2a1 (Q-L54)
ARS006 F C4a1a+195 -
ARS007 M A+152+16362 Q1a2a (Q-L475; Q-L53)
ARS008 M C4a2c Q1a2a1c (Q-L330)
ARS009 F D5a2a -
ARS011 M n/d Q1a2 (Q-L56; Q-M346)
ARS012 F C4a2a1 -
ARS013 F C -
ARS014 F C4a2c1 -
ARS015 M G3a Q1a1 (Q-Y706; Q-F1096)
ARS016 M A+152+16362+16189 Q1a2a1c (Q-L334; Q-L330)
ARS017 F G2a -
ARS018 M B5b1 Q1a (Q-M1117; Q-L472)
ARS024 F C4a1a1 -
ARS025 M D4b1a2a Q1a2a (Q-L475; Q-L53)
ARS026 M C4a1a+195 R1a1a1b2a2a (R-Z2123)
AST001 F I1a -
ATS001 F C4a1a -
BAA001 M D4j5a C2b (C-F2661; C-L1373)
BAM001 F D4j3 -
BAU001 F N1a1a1a1 -
BAY001 M D4e5b C2b1a1b (C-F3937; C-F1756)
BAZ001 F G3a3 -
BER002 M T1a1 N1c1 (N-L395; N-M46)
BIL001 F F1b1b -
BOR001 F - -
BRG001 F M7b1a1a2 -
BRG002 F G2a5 -
BRG004 M D4c2a CT (CT-M5613; CT-M168)
BRG005 M G1a N1c2b2 (N-L665)
BRL001 F G1a1a -
BRL002 F HV6 -
BRL003 M - C2b1a1b (C-F1756)
BRU001 M D5b1b2 J1a2b (J-P58)
BTO001 M Z4a Q1a2a1c (Q-L330)
BUL001 M M11b Q1a1a1 (Q-M120)
BUL002 M M10a1b G2a2b2a1a1c1a2a1a (G-FGC226; G-FGC249)
BUR001 F T2b -
BUR002 M U2e1 E1b1b1a1b2 (E-V22; E-L677)
BUR003 M J2b1a2a R1a1a1b (R-Z647; R-Z645)
BUR004 F G2a5 -
BUU001 F C4a1a4a -
CHD001 F F1g -
CHN001 M T1a1 R1a1a1b2a2b (R-Z2122)
CHN003 M U4a1 Q1a2a (Q-L213; Q-L53)
CHN004 F H91 -
CHN006 F U2e1h -
CHN007 M U2e1h Q1a2a1c1 (Q-L332; Q-L329)
CHN008 M F1b1f R1a1a1b2 (R-Z93)
CHN010 M D4j12 C2b1a1b (C-F1756)
CHN012 M T2b34 R1a1a1b (R-Z645)
CHN013 F D4g1b -
CHN014 F D4l2 -
CHN015 F N9a9 -
CHN016 M A8a Q1a2a1c1 (Q-L332; Q-L329)
DAR001 M F1b1f Q1b2 (Q-Y1151; Q-Y1150)
DAR002 F A11 -
DAS001 M N1a1a1a1a C2b1c (C-M546; C-F1918)
DEE001 F B4d1'2'3 -
DEK001 / SHR001 M R11a O2a2b1a1a3a (O-F474; O-F317)
DEL001 M D4j11 Q1a2a1c1 (Q-L332; Q-L329)
DOL001 F C4a1a -
DUU001 F D4b1a2a1 -
DUU002 M F2a R1a1a1 (R-Page7; R-M417)
EME002 F B5a2a1b -
EME003 M D5b1b2 O2a2b1a2a1 (O-F388; O-F46)
ERD001 F B4f -
ERM001 / ERM002 / ERM003 F A12 -
FNO001 M C4 N
FNO003 F C -
FNO006 F C4a1a3 -
FNO007 F F1b1+@152 -
GAN002 F A11a -
GTO001 F M7b1b -
GUN002 F F1a1 -
GUN003 M D5b1 N1 (N-L735)
HUD001 F K1a19 -
IAG001 M T2b2 R1b1a2 (R-CTS12478; R-M269)
IKU001 F F1b1b -
IMA001 F C4a1a+195 -
IMA002 F D4j5a -
IMA003 M D4j5a R1a1
IMA004 M G2a+152 Q1a2a
IMA005 M D4j5a N1c1a1a
IMA006 M A24 Q1a1a1
IMA007 F A+152+16362 -
IMA008 F N9a -
JAA001 F M10a1b -
JAG001 M F1b1f -
KGK001 F D4b1a2a1 -
KHI001 M R11 N1c2b2 (N-L665)
KHL001 F F1b1b -
KHN001 / KHN002 M D4j5a C2b1a1b1 (C-F3830)
KHO001 M D4 J (J-CTS10446; J-M304)
KHO003 F D4j5 -
KHO006 M C5b1b C2b1a1b (C-F3937; C-F1756)
KHO007 M - CT (CT-M5603; CT-M168)
KHU001 M C4a1a Q1a2a (Q-L55; Q-L53)
KHV001 M B4b1a3a CT (CT-M5606; CT-M168)
KHV002 M C5a1 N1c2b2 (N-L665)
KNN001 M Z3c O2a1c1a5 (O-M5420)
KNU001 F F1b1f -
KRN001 M Z3c C2b1b1 (C-M86)
KRN002 F A18 -
KUM001 F A -
KUR001 F A -
MIT001 M D1j Q1a1a1 (Q-M120)
MRI001 M D4m D (D-F1137; D-M174)
NAI001 M K1a12a1a J2a1h2 (J-L25)
NAI002 / NAI003 M C4a2c1 R1a1a1b1 (R-Z283)
NOM001 M U7a3a R1a1a1b2 (R-Z93)
NRC001 F A -
OLN001.A F D4 -
OLN001.B M B5b1a Q1a1 (Q-F1215; Q-F1096)
OLN002 F T1a1 -
OLN003 M T1a1 R1b1a2 (R-PF6505; R-M269)
OLN004 F H4a1a1a -
OLN005 M H6b R1b1a2a2 (R-Z2105; R-CTS1078)
OLN007 M D4 C2e1a1a (C-F3864; C-M407)
OLN008 F D4i -
OLN009 F J1b1b1 -
OLN010 M G3a3 R1b1a2a2 (R-Z2105; R-CTS1078)
OLN011 M B5b2a R1b1a2a2 (R-CTS1078)
OLN012 F D4j10 -
PTO001 F D4j+(16286) -
RAH001 F F1b1e -
SAN001 F F1b1e -
SBG001 M G2b2 Q1a2a1 (Q-L54)
SHA001 M D4a1 C2b1c (C-F3795; C-F1918)
SHG001 M M9a1a C2b1a1b1 (C-F3830)
SHG002 M D4o1 N1c1a1 (N-L708)
SHG003 M G2a2a C2b1c (C-F4002; C-F1918)
SHT001 M N1a1a1a1 R1b1a2a1a (R-P311)
SHT002 M U5a1 J1a (J-CTS5368)
SHU001 M G2b2a Q1a1a1 (Q-M120)
SHU002 M D4b2b R1a1a1b (R-Z645)
SKT001 F U5a1i -
SKT002 M G2a1e R1b (R-M343)
SKT003 F I1a -
SKT004 F U2e1a1 -
SKT005 M F1b1f R1b1 (R-M415; R-P25_1)
SKT006 M G2a1e R1 (R-P236; R-M173)
SKT007 M C4 Q1a2a1c (Q-L334; Q-L330)
SKT008 M H Q1a2a1c (Q-L334; Q-L330)
SKT009 M I1a R1a1a1b (R-Z647; R-Z645)
SKT010 F I1a -
SKT011 F D5a2a1 -
SKT012 M C4a1a4a J2a1h2 (J-L25)
SOL001 F I4a -
SON001 M D4 NO (NO-M2308; NO-M2313)
SOU001 F F1b1 -
TAH002 M B4b1b D1a2a (D-P47)
TAK001 F G2a1 -
TAV001 M M9a R1a1a1b (R-Z645)
TAV005 M D4j+16311 R1b1a2(R-PF6475,R-M269)
TAV006 F HV2a2 -
TAV007 M D4j3 R1b1a2a1a1 (R-M405)
TAV011 M C4b1 R1a1a1b2a2a (R-Z2123)
TEV002 F C5b -
TEV003 M N9a1 C2b1a1b(C-F1756)
TMI001 F J2b1a2a -
TSA001 M T1a1 C2b1a1b1 (C-F3830)
TSA002 M D4o2a C2b1 (C-F1788; C-F1699)
TSA003 M D4j3 C2b1c (C-F3939; C-F1918)
TSA004 M D4a1 D1a1a (D-N1)
TSA005 M Z1a N1c1 (N-L395; N-M46)
TSA006 F F1b1f -
TSA007 F M8a2'3 -
TSB001 M H14b J1a (J-CTS5368)
TSI001 F F1b1f -
TUH001 M B5a2a1a C2b1b1 (C-M86)
TUH002 M B5a2a1a J2a (J-M410)
TUK001/TAV008 F H2a2b -
TUK002 M B5b2a2 O2a1c1a5 (O-M5420)
TUK003 F J1d5 -
TUM001 M F1c1a1 N1c2 (N-L666)
UAA001 M H15b1 NO1 (NO-M214)
UGO001 F K2a5b -
UGO002 M U5a1a1 O2a2b1a1 (O-Page23)
UGU001 M D2 J2a (J-L559; J-M410)
UGU002 M B4a2b1 R (R-P224; R-M207)
UGU003 F U3a -
UGU004 F C4a1a4a -
UGU005 M U4a1 R1a1a1b2a2a (R-Z2123)
UGU006 M Z1a J1 (J-M267)
UGU010 M - R1a1a1b2a2a (R-Z2123)
UGU011 F HV6 -
ULA001 M Y1a J2a2 (J-L581)
ULI001 M U2e1 R1a1a1b2a2b (R-Z2122)
ULI002 M B4c1a2a J2a (J-L559; J-M410)
ULI003 F H6a1b2 -
ULN001 M D5a2a1b Q1a1a1 (Q-M265; Q-M120)
ULN002 F C4a1b -
ULN003 F C4a1a -
ULN004 F C5a1 -
ULN005 F D4 -
ULN006 F C4 -
ULN007 M D4b1a2 Q1a1 (Q-F1096)
ULN009 F C4a1a -
ULN010 F D5a2a -
ULN011 M A2a3 O2a2b1a1a (O-F42; O-F8)
ULN015 F D5a2a1b -
ULZ001 F D4b1a2 -
UUS001 M D5a2a J1a2b (J-P58)
UUS002 M X2d1 Q1a1a1 (Q-M265; Q-M120)
UVG001 F G2a2 -
YAG001 M T1a1 R1b1a2a2 (R-CTS1078)
YUR001 M D5a2a1 N1 (N-L735)
ZAA001 F A24 -
ZAA002 M C4a2a1 C2b1a1b1 (C-F3830)
ZAA003 F F2a -
ZAA004 M D4b1a2a1 C2b1a1b1 (C-F3830)
ZAA005 M D4c1b1 J2a (J-L212; J-M410)
ZAA007 F D4l2 -
ZAM001 M D4q R1a1a (R-L449; R-M512)
ZAM002 M B6a D1a1a (D-N1)
ZAR002 F F1 -
ZAY001 M M9a1b2 O2a2b1a1a (O-F8)

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