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Cited by 331Forensic and Genetic ResearchGenetic diversity and population structureEurasian Exchange Networks

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

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Jeong, Choongwon, Wang, Ke, Wilkin, Shevan, Taylor, William Timothy Treal, Miller, Bryan K., Bemmann, Jan H., Stahl, Raphaela, Chiovelli, Chelsea, Knolle, Florian, Ulziibayar, Sodnom, Khatanbaatar, Dorjpurev, Erdenebaatar, Diimaajav, Erdenebat, Ulambayar, Ochir, Ayudai, Ankhsanaa, Ganbold, Vanchigdash, Chuluunkhuu, Ochir, Battuga, Munkhbayar, Chuluunbat, Tumen, Dashzeveg, Kovalev, Alexey, Kradin, Nikolay, Bazarov, Bilikto A., Miyagashev, Denis A., Konovalov, Prokopiy B., Zhambaltarova, Elena, Miller, Alicia Ventresca, Haak, Wolfgang, Schiffels, Stephan, Krause, Johannes, Boivin, Nicole, Erdene, Myagmar, Hendy, Jessica, Warinner, Christina·2020·Cell
DOI 10.1016/j.cell.2020.10.015PMC PMC7664836

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Abstract

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.

Overview

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TL;DR

This paper reports genome-wide data for 214 ancient individuals from Mongolia and neighboring Russia spanning ~6,000 years, revealing a dynamic population history of the Eastern Steppe. It documents a pastoralist expansion ca. 3000 BCE, a tripartite Bronze Age genetic structure, Xiongnu-era admixture, and a Mongol-period shift toward higher eastern Eurasian ancestry.

Key findings

  • Pre-Bronze Age hunter-gatherers carried Ancient Northeast Asian (ANA) ancestry; Afanasievo individuals in central Mongolia (3112–2917 cal. BCE) were genetically indistinguishable from Yenisei Afanasievo, confirming Western Steppe herder expansion ~1,500 km east, but Afanasievo and Chemurchek left little enduring genetic trace in later MLBA populations.
  • By the Middle/Late Bronze Age (1900–900 BCE), Mongolia was biogeographically structured into three distinct gene pools: Khövsgöl_LBA, Altai_MLBA (admixed between Khövsgöl_LBA and Sintashta-related Western Steppe ancestry), and Ulaanzuukh_SlabGrave (ANA-related, lacking ANE and WSH admixture); all practiced dairy pastoralism regardless of ancestry.
  • The Xiongnu emerged from mixing these populations and surrounding regions: earlyXiongnu_west derived 92% ancestry from Chandman_IA with additional Iranian-related ancestry, while late Xiongnu showed added Sarmatian-related and Han-related gene flow and extreme genetic heterogeneity.
  • Mongol-era individuals had much higher eastern Eurasian ancestry, modeled as 55%–64% Ulaanzuukh_SlabGrave, 21%–27% Han-related, and 15%–18% Western Steppe ancestry; 34 of 61 historic Mongols were genetically cladal with at least one modern Mongolic-speaking population, and lactase persistence remained at negligibly low frequency (~5%) despite widespread dairy consumption.

Methods

  • New genome-wide data were generated for 214 ancient individuals from 85 Mongolian and 3 Russian sites, plus published data; 30 new AMS radiocarbon dates supplemented 74 previously published dates, totaling 98 directly dated individuals (104 dates).
  • Sampling included 169 teeth and 75 petrosal bones from 225 individuals; 24 individuals were excluded for insufficient human DNA or <10,000 SNPs after capture, and 10 for >5% contamination or uncertain genetic sex; genetic sex was determined for 224 individuals (100 female, 124 male).
  • Analyses included PCA projected onto 2,077 present-day Eurasians, outgroup f3 and f4 statistics, qpAdm/qpWave admixture modeling with eight outgroups, DATES admixture dating assuming 29 years per generation, uniparental haplogroup assignment, relatedness estimation, and allele-frequency estimation for five phenotypic SNPs.

Limitations

  • The paper states that Early Medieval sampling is uneven (1 Xianbei/Rouran, 8 Türkic, 13 Uyghur individuals), Khitan representation is small (n = 3), and Mongol-era sampling excludes royal/regional elites and Karakorum; it also calls for further research in central and eastern Eurasia, northeastern China, the Tarim Basin, and the eastern Kazakh steppe.
  • The paper states that some DATES admixture estimates should be interpreted with care because the method is designed for 2-way admixture but applied to 3-way models, and that Chemurchek source populations are distant in time or space; it also notes that ancient China, Korea, Japan, and Southeast Asia remain mostly unsampled.
  • Evident from the text: nomadic camp sites are ephemeral and poorly preserved, knowledge of Mongolian prehistory is biased toward the past five millennia, radiocarbon dating reassigned many burials across periods, and sex-biased and kinship inferences are limited by sample size and context.

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Crossref · 10.1016/j.cell.2020.10.015

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Figures

Graphical abstract
Graphical abstract CC BY · PubMed Central
Figure 1 Overview of Ancient Populations and Time Periods (A) Distribution of sites with their associated culture and time period indicated by color: Pre-Bronze, purple; Early Bronze, red; Middle/Late Bronze, blue; Early Iron, pink; Xiongnu, green; Early Medieval, brown; Late Medieval, gold (see STAR Methods). See Figure S1A and Table S1B for site codes and labels. (B) Inset map of Eurasia indicating area of present study (box) and the locations of other ancient populations referenced in the text, colored by time period. The geographic extent of the Western/Central Steppe is indicated in light brown, and the Eastern Steppe is indicated in light green. (C) Timeline of major temporal periods and archaeological cultures in Mongolia. Site locations have been jittered to improve visibility of overlapping sites.
Figure 1 Overview of Ancient Populations and Time Periods (A) Distribution of sites with their associated culture and time period indicated by color: Pre-Bronze, purple; Early Bronze, red; Middle/Late Bronze, blue; Early Iron, pink; Xiongnu, green; Early Medieval, brown; Late Medieval, gold (see STAR Methods). See Figure S1A and Table S1B for site codes and labels. (B) Inset map of Eurasia indicating area of present study (box) and the locations of other ancient populations referenced in the text, colored by time period. The geographic extent of the Western/Central Steppe is indicated in light brown, and the Eastern Steppe is indicated in light green. (C) Timeline of major temporal periods and archaeological cultures in Mongolia. Site locations have been jittered to improve visibility of overlapping sites.CC BY · PubMed Central
Figure S1 Archaeological Sites and Geographic and Ecological Features in Mongolia, Related to Figure 1 (A) Archaeological sites in Mongolia and neighboring regions analyzed in this study. (B) Mongolian regions and provinces (aimags). Provinces are indicated by gray lines and text. Regions are indicated by black dashed lines and text following the definitions of (Taylor et al., 2019). (C) Ecological zones of Mongolia. Map produced using QGIS software (v3.6) with ecological data from (Dorjgotiv, 2004).
Figure S1 Archaeological Sites and Geographic and Ecological Features in Mongolia, Related to Figure 1 (A) Archaeological sites in Mongolia and neighboring regions analyzed in this study. (B) Mongolian regions and provinces (aimags). Provinces are indicated by gray lines and text. Regions are indicated by black dashed lines and text following the definitions of (Taylor et al., 2019). (C) Ecological zones of Mongolia. Map produced using QGIS software (v3.6) with ecological data from (Dorjgotiv, 2004).CC BY · PubMed Central
Figure 2 Genetic Structure of Mongolia through Time PCA of ancient individuals (n = 214) from three major periods projected onto contemporary Eurasians (gray symbols). Main panels display PC1 versus PC2; insets display PC1 versus PC3. Inset tick marks for PC1 correspond to those for the main panels; PC3 accounts for 0.35% of variation. See Figure S3B for population, sample, and axis labels, and Tables S1B, S1C, and S2A for further site and sample details.
Figure 2 Genetic Structure of Mongolia through Time PCA of ancient individuals (n = 214) from three major periods projected onto contemporary Eurasians (gray symbols). Main panels display PC1 versus PC2; insets display PC1 versus PC3. Inset tick marks for PC1 correspond to those for the main panels; PC3 accounts for 0.35% of variation. See Figure S3B for population, sample, and axis labels, and Tables S1B, S1C, and S2A for further site and sample details.CC BY · PubMed Central
Figure 3 Genetic Changes in the Eastern Steppe across Time Characterized by qpAdm (A–F) Major time periods: (A) Pre-Bronze through Early Bronze Age, (B) Middle/Late Bronze Age, (C) Early Iron Age, (D) Xiongnu period, (E) Early Medieval, and (F) Late Medieval. Modeled ancestry proportions are indicated by sample size-scaled pie charts, with ancestry source populations shown below (see STAR Methods). The sample size range for each panel is indicated in the upper right. For (B) and (C), Baikal_EBA is modeled as light blue; in (D–F), Khövsgöl_LBA (purple) and the Sagly/Uyuk of Chandman_IA (pink) are modeled as new sources (Figure 4). Cultural groups are indicated by bold text. For (D–F), individuals are Late Xiongnu, Türkic, and Mongol, respectively, unless otherwise noted. Previously published reference populations are noted with white text; all others are from this study. Populations beyond the map borders are indicated by arrows. Burial locations have been jittered to improve visibility of overlapping individuals.
Figure 3 Genetic Changes in the Eastern Steppe across Time Characterized by qpAdm (A–F) Major time periods: (A) Pre-Bronze through Early Bronze Age, (B) Middle/Late Bronze Age, (C) Early Iron Age, (D) Xiongnu period, (E) Early Medieval, and (F) Late Medieval. Modeled ancestry proportions are indicated by sample size-scaled pie charts, with ancestry source populations shown below (see STAR Methods). The sample size range for each panel is indicated in the upper right. For (B) and (C), Baikal_EBA is modeled as light blue; in (D–F), Khövsgöl_LBA (purple) and the Sagly/Uyuk of Chandman_IA (pink) are modeled as new sources (Figure 4). Cultural groups are indicated by bold text. For (D–F), individuals are Late Xiongnu, Türkic, and Mongol, respectively, unless otherwise noted. Previously published reference populations are noted with white text; all others are from this study. Populations beyond the map borders are indicated by arrows. Burial locations have been jittered to improve visibility of overlapping individuals.CC BY · PubMed Central
Figure S2 Uniparental Haplogroup Assignments by Group and Sex-Bias Z Scores, Related to Figure 5B and STAR Methods (A and B) Population structure from uniparentally inherited markers. (A) Distribution of Y haplogroups across each period. (B) Distribution of mitochondrial haplogroups across each period. (C) Sex-bias Z scores by evaluating the differences of WSH-/Iranian-/Han-related ancestry on the autosomes and the X chromosome. We calculated Z-score for each ancient individual who has genetic admixture with any of the three ancestries. Positive scores suggest more WSH-/Iranian-/Han-related ancestry on the autosomes, i.e., male-driven admixture.
Figure S2 Uniparental Haplogroup Assignments by Group and Sex-Bias Z Scores, Related to Figure 5B and STAR Methods (A and B) Population structure from uniparentally inherited markers. (A) Distribution of Y haplogroups across each period. (B) Distribution of mitochondrial haplogroups across each period. (C) Sex-bias Z scores by evaluating the differences of WSH-/Iranian-/Han-related ancestry on the autosomes and the X chromosome. We calculated Z-score for each ancient individual who has genetic admixture with any of the three ancestries. Positive scores suggest more WSH-/Iranian-/Han-related ancestry on the autosomes, i.e., male-driven admixture.CC BY · PubMed Central
Figure S3 PCA of Present-Day Eurasian Populations and Genetic Structure of Mongolia through Time, Related to Figure 2 (A) PCA of present-day Eurasian populations used as the background for Figure 2 and Figure S3B. Here we show the population labels for the 2,077 Eurasian individuals used for calculating PCs and plotted as gray dots in Figure 2. Each three-letter code in the plot represents a single individual. Population IDs matching to the three-letter codes are listed at the bottom. (B) Genetic structure of Mongolia through time. Principal component analysis (PCA) of ancient individuals (n = 214) from three major periods projected onto contemporary Eurasians (gray symbols). Projection and axis variance corresponds to Figure 2. Population labels are positioned over the mean coordinate across individuals belonging to each population.
Figure S3 PCA of Present-Day Eurasian Populations and Genetic Structure of Mongolia through Time, Related to Figure 2 (A) PCA of present-day Eurasian populations used as the background for Figure 2 and Figure S3B. Here we show the population labels for the 2,077 Eurasian individuals used for calculating PCs and plotted as gray dots in Figure 2. Each three-letter code in the plot represents a single individual. Population IDs matching to the three-letter codes are listed at the bottom. (B) Genetic structure of Mongolia through time. Principal component analysis (PCA) of ancient individuals (n = 214) from three major periods projected onto contemporary Eurasians (gray symbols). Projection and axis variance corresponds to Figure 2. Population labels are positioned over the mean coordinate across individuals belonging to each population.CC BY · PubMed Central
Figure 4 Genetic Ancestry Changes in Chronological Order across All Newly Reported Genetic Groups Well-fitted modeling results for grouped-based population genetics analyses for (A) prehistoric periods and (B) historic periods. The number of individuals in each genetic group is given in Table S3A. Raw ancestry proportions and standard error estimates are provided in Table S5. Horizontal bars represent ± 1 standard error (SE) estimated by qpAdm.
Figure 4 Genetic Ancestry Changes in Chronological Order across All Newly Reported Genetic Groups Well-fitted modeling results for grouped-based population genetics analyses for (A) prehistoric periods and (B) historic periods. The number of individuals in each genetic group is given in Table S3A. Raw ancestry proportions and standard error estimates are provided in Table S5. Horizontal bars represent ± 1 standard error (SE) estimated by qpAdm.CC BY · PubMed Central
Figure S4 Genetic Changes in the Eastern Steppe across Time Characterized by qpAdm with All Individuals Indicated, Related to Figures 3 and 4 (A) Pre-Bronze through Early Bronze Age; (B) Middle/Late Bronze Age; (C) Early Iron Age; (D) Xiongnu period; (E) Early Medieval; (F) Late Medieval. Modeled ancestry proportions are indicated by sample size-scaled pie charts, with ancestry source populations shown below. Cultural groups are indicated by bold text. For panels (D–F), individuals are Late Xiongnu, Türkic, and Mongol, respectively, unless otherwise noted. Previously published reference populations are noted with white text; all others are from this study. Populations beyond the map borders are indicated by arrows. Burial locations have been jittered to improve visibility of overlapping individuals. Zoom in to see individual labels. Here we report results from admixture models that include all ancestry components required to explain historic late Medieval individuals as a group for unbiased cross comparison between individuals. Individual results with simpler admixture models can be found in Table S5J. See modeling details in Section 7.
Figure S4 Genetic Changes in the Eastern Steppe across Time Characterized by qpAdm with All Individuals Indicated, Related to Figures 3 and 4 (A) Pre-Bronze through Early Bronze Age; (B) Middle/Late Bronze Age; (C) Early Iron Age; (D) Xiongnu period; (E) Early Medieval; (F) Late Medieval. Modeled ancestry proportions are indicated by sample size-scaled pie charts, with ancestry source populations shown below. Cultural groups are indicated by bold text. For panels (D–F), individuals are Late Xiongnu, Türkic, and Mongol, respectively, unless otherwise noted. Previously published reference populations are noted with white text; all others are from this study. Populations beyond the map borders are indicated by arrows. Burial locations have been jittered to improve visibility of overlapping individuals. Zoom in to see individual labels. Here we report results from admixture models that include all ancestry components required to explain historic late Medieval individuals as a group for unbiased cross comparison between individuals. Individual results with simpler admixture models can be found in Table S5J. See modeling details in Section 7.CC BY · PubMed Central
Figure S5 Outgroup f3-Statistics and Cladality Testing using f4-Statistics, Related to Figures 3 and 4 (A) Outgroup f3-statistics for the pre-Bronze Age to Early Bronze Age groups in the Eastern Steppe. We show top 15 outgroup f3-statistics of the form f3(Target, world-wide; Mbuti) out of 345 ancient and present-day populations for the six target groups: eastMongolia_preBA, centralMongolia_preBA, Fofonovo_EN, Afanasievo_Mongolia, Chemurchek_southAltai and Chemurchek_northAltai. Horizontal bars represent ± 1 standard error (SE) calculated by 5 cM block jackknifing. (B) Testing cladality of the four ANA populations using f4-statistics. We show top and bottom 15 symmetric f4-statistics of the form f4(ANA1, ANA2; world-wide, Mbuti) out of 345 ancient and present-day populations for the four ANA-related target groups: eastMongolia_preBA, centralMongolia_preBA, Fofonovo_EN, DevilsCave_N. Horizontal bars represent ± 1 standard error (SE) calculated by 5 cM block jackknifing. f4-statistics with Z-score > 3 are highlighted in red. (C) Testing cladality of Afanasievo and Chemurchek using f4-statistics. We show top and bottom 15 symmetric f4-statistics for the three target groups Afanasievo_Mongolia, Chemurchek_southAltai and Chemurchek_northAltai, in the form f4(Afanasievo_Mongolia, Afanasievo; world-wide, Mbuti), f4(Chemurchek_southAltai, Botai; world-wide, Mbuti), f4(Chemurchek _northAltai, Chemurchek_southAltai; world-wide, Mbuti), and f4(Chemurchek _northAltai, eastMongolia_preBA; world-wide, Mbuti) out of 345 ancient and present-day populations. Horizontal bars represent ± 1 standard error (SE) calculated by 5 cM block jackknifing. f4-statistics with Z-score > 3 are highlighted in red.
Figure S5 Outgroup f3-Statistics and Cladality Testing using f4-Statistics, Related to Figures 3 and 4 (A) Outgroup f3-statistics for the pre-Bronze Age to Early Bronze Age groups in the Eastern Steppe. We show top 15 outgroup f3-statistics of the form f3(Target, world-wide; Mbuti) out of 345 ancient and present-day populations for the six target groups: eastMongolia_preBA, centralMongolia_preBA, Fofonovo_EN, Afanasievo_Mongolia, Chemurchek_southAltai and Chemurchek_northAltai. Horizontal bars represent ± 1 standard error (SE) calculated by 5 cM block jackknifing. (B) Testing cladality of the four ANA populations using f4-statistics. We show top and bottom 15 symmetric f4-statistics of the form f4(ANA1, ANA2; world-wide, Mbuti) out of 345 ancient and present-day populations for the four ANA-related target groups: eastMongolia_preBA, centralMongolia_preBA, Fofonovo_EN, DevilsCave_N. Horizontal bars represent ± 1 standard error (SE) calculated by 5 cM block jackknifing. f4-statistics with Z-score > 3 are highlighted in red. (C) Testing cladality of Afanasievo and Chemurchek using f4-statistics. We show top and bottom 15 symmetric f4-statistics for the three target groups Afanasievo_Mongolia, Chemurchek_southAltai and Chemurchek_northAltai, in the form f4(Afanasievo_Mongolia, Afanasievo; world-wide, Mbuti), f4(Chemurchek_southAltai, Botai; world-wide, Mbuti), f4(Chemurchek _northAltai, Chemurchek_southAltai; world-wide, Mbuti), and f4(Chemurchek _northAltai, eastMongolia_preBA; world-wide, Mbuti) out of 345 ancient and present-day populations. Horizontal bars represent ± 1 standard error (SE) calculated by 5 cM block jackknifing. f4-statistics with Z-score > 3 are highlighted in red.CC BY · PubMed Central
Figure S6 Dating Admixture in Prehistoric Individuals, Related to STAR Methods (A) Dating admixture in prehistoric individuals. We estimated admixture dates using the DATES program and converted it by adding the age of each ancient population (mean value of the center of the 95% confidence interval of calibrated 14C dates) and assuming 29 years per generation. Horizontal bars associated with the admixture dates (colored circles) are estimated by the square root of summing the variance of DATES estimate using leave-one-chromosome-out jackknifing method and the variance of the 14C date estimate, assuming that the two quantities are independent. Published groups are marked with an asterisk (∗). For the Chemurchek_northAltai, we used Baikal_EN as the representative of ANA ancestry for dating the admixture event, given the larger sample size of Baikal_EN. (B) Ancestry covariance in prehistoric individuals. We show the weighted ancestry covariance (y axis) calculated from DATES which is expected to decay exponentially along genetic distance (x axis) with a decay rate indicating the time since admixture, and fitted exponential curves (shown in red line). We start the fit at genetic distance at 0.45 centiMorgans, and estimate standard error by a weighted block jackknife removing one chromosome in each run.
Figure S6 Dating Admixture in Prehistoric Individuals, Related to STAR Methods (A) Dating admixture in prehistoric individuals. We estimated admixture dates using the DATES program and converted it by adding the age of each ancient population (mean value of the center of the 95% confidence interval of calibrated 14C dates) and assuming 29 years per generation. Horizontal bars associated with the admixture dates (colored circles) are estimated by the square root of summing the variance of DATES estimate using leave-one-chromosome-out jackknifing method and the variance of the 14C date estimate, assuming that the two quantities are independent. Published groups are marked with an asterisk (∗). For the Chemurchek_northAltai, we used Baikal_EN as the representative of ANA ancestry for dating the admixture event, given the larger sample size of Baikal_EN. (B) Ancestry covariance in prehistoric individuals. We show the weighted ancestry covariance (y axis) calculated from DATES which is expected to decay exponentially along genetic distance (x axis) with a decay rate indicating the time since admixture, and fitted exponential curves (shown in red line). We start the fit at genetic distance at 0.45 centiMorgans, and estimate standard error by a weighted block jackknife removing one chromosome in each run.CC BY · PubMed Central
Figure S7 Breakdown of Geography and Genetics among Xiongnu and Comparison of Mongol Period and Present-Day Populations, Related to Figure 3 and STAR Methods (A) Breakdown of the geographic-genetic correlation in Xiongnu. We show the proportions of West Eurasian ancestry on all individuals/groups from Xiongnu era (y axis) versus the longitude of archaeological site they come from (x axis). The raw numbers of individual estimates can be found in Table S5G for models using Sarmatian as the western Eurasian source. Unlike MLBA/EIA individuals (Figure 3), Xiongnu individuals from more western sites do not have higher proportion of western Eurasian ancestry than those from eastern sites. (B) Comparing genetic homogeneity between ancient Mongol individuals and seven present-day Mongolic-speaking populations using qpWave. We report the p-value for every individual-based qpWave {ancient Mongol individual; Mongolic group} using seven modern Mongolic-speaking populations: Buryat, Daur, Kalmyk, Khamnegan, Mongol, Mongola, and Tu in the Human Origins dataset. When the p-value from qpWave is > 0.05, it suggests that the ancient individual on the y axis is genetically indistinguishable from the modern Mongolic-speaking population shown on the x axis. Smaller p-values indicate that the ancient individual is significantly different from the modern group.
Figure S7 Breakdown of Geography and Genetics among Xiongnu and Comparison of Mongol Period and Present-Day Populations, Related to Figure 3 and STAR Methods (A) Breakdown of the geographic-genetic correlation in Xiongnu. We show the proportions of West Eurasian ancestry on all individuals/groups from Xiongnu era (y axis) versus the longitude of archaeological site they come from (x axis). The raw numbers of individual estimates can be found in Table S5G for models using Sarmatian as the western Eurasian source. Unlike MLBA/EIA individuals (Figure 3), Xiongnu individuals from more western sites do not have higher proportion of western Eurasian ancestry than those from eastern sites. (B) Comparing genetic homogeneity between ancient Mongol individuals and seven present-day Mongolic-speaking populations using qpWave. We report the p-value for every individual-based qpWave {ancient Mongol individual; Mongolic group} using seven modern Mongolic-speaking populations: Buryat, Daur, Kalmyk, Khamnegan, Mongol, Mongola, and Tu in the Human Origins dataset. When the p-value from qpWave is > 0.05, it suggests that the ancient individual on the y axis is genetically indistinguishable from the modern Mongolic-speaking population shown on the x axis. Smaller p-values indicate that the ancient individual is significantly different from the modern group.CC BY · PubMed Central
Figure 5 Functional Allele Frequencies and Sex-Biased Patterns of Genetic Admixture (A) Allele frequencies of five phenotypic SNP changes through time. For the effective allele, we show maximum likelihood frequency estimates and one standard error bar for each ancient group. The pre-MLBA category corresponds to the sum of all ancient groups before Mönkhkhairkhan. Xiongnu, Early Medieval, and Late Medieval correspond to the sum of all ancient groups in each period correspondingly. Horizontal dashed lines show allele frequency information from the 1000 Genomes Project’s five super populations. (B) Sex-biased patterns of genetic admixture by period and population. We calculated Z scores for every ancient individual who has genetic admixture with WSH-/Iranian-/Han-related ancestry. Positive scores suggest more WSH-/Iranian-/Han-related ancestry on the autosomes, i.e., male-driven admixture. See Figure S2C for individual Z scores.
Figure 5 Functional Allele Frequencies and Sex-Biased Patterns of Genetic Admixture (A) Allele frequencies of five phenotypic SNP changes through time. For the effective allele, we show maximum likelihood frequency estimates and one standard error bar for each ancient group. The pre-MLBA category corresponds to the sum of all ancient groups before Mönkhkhairkhan. Xiongnu, Early Medieval, and Late Medieval correspond to the sum of all ancient groups in each period correspondingly. Horizontal dashed lines show allele frequency information from the 1000 Genomes Project’s five super populations. (B) Sex-biased patterns of genetic admixture by period and population. We calculated Z scores for every ancient individual who has genetic admixture with WSH-/Iranian-/Han-related ancestry. Positive scores suggest more WSH-/Iranian-/Han-related ancestry on the autosomes, i.e., male-driven admixture. See Figure S2C for individual Z scores.CC BY · PubMed Central

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Authors
Jeong, C., Wang, K., Wilkin, S., Taylor, W. T. T., Miller, B. K., Bemmann, J. H., Stahl, R., Chiovelli, C., Knolle, F., Ulziibayar, S., Khatanbaatar, D., Erdenebaatar, D., Erdenebat, U., Ochir, A., Ankhsanaa, G., Vanchigdash, C., Ochir, B., Munkhbayar, C., Tumen, D., Kovalev, A., Kradin, N., Bazarov, B. A., Miyagashev, D. A., Konovalov, P. B., Zhambaltarova, E., Miller, A. V., Haak, W., Schiffels, S., Krause, J., Boivin, N., Erdene, M., Hendy, J., & Warinner, C.
Affiliations
  • Seoul National University (KR)
  • Max Planck Institute of Geoanthropology (DE)
  • University of Colorado Boulder (US)
  • University of Michigan (US)
  • University of Bonn (DE)
  • Mongolian Academy of Sciences (MN)
  • Mongolian University of Science and Technology (MN)
  • National University of Mongolia (MN)
  • Russian Academy of Sciences (RU)
  • East Siberian Academy of Culture and Arts (RU)
  • Harvard University (US)
Corresponding author
  • Jeong, Choongwon
  • Warinner, Christina
ORCID
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Cell
Publisher
Elsevier BV
Published
2020-11
Volume
183
Issue
4
Pages
890-904.e29
ISSN
0092-8674 (ISSN-L)
DOI
10.1016/j.cell.2020.10.015
PMC
PMC7664836
OpenAlex
W3094811729
Cited by
328
References
113
Citations by year
2026: 59 · 2025: 88 · 2024: 51 · 2023: 28 · 2022: 48 · 2021: 48 · 2020: 6
FWCI
15.68
Type
journal article
Language
en
Open access
Licence
Locations
  • Cell (publishedVersion, cc-by)
  • bioRxiv (Cold Spring Harbor Laboratory) (submittedVersion, cc-by-nc-nd)
  • PubMed (publishedVersion)
  • PubMed Central (publishedVersion, cc-by)
  • White Rose Research Online (University of Leeds, The University of Sheffield, University of York) (acceptedVersion)
  • MPG.PuRe (Max Planck Society) (submittedVersion, other-oa)
  • Griffith Research Online (submittedVersion, cc-by)
  • Seoul National University Open Repository (Seoul National University) (submittedVersion)
  • Cell
Funders
  • Deutsche Forschungsgemeinschaft (257731206, SFB 1167)
  • Russian Foundation for Basic Research (18-59-94020)
  • Ministry of Science, ICT and Future Planning (2020R1C1C1003879)
  • Ministry of Education and Science of the Russian Federation
  • National University of Mongolia
  • Horizon 2020
  • National Research Foundation of Korea
  • Max-Planck-Gesellschaft
  • Horizon 2020 Framework Programme
  • European Research Council (804884-DAIRYCULTURES, 771234-PALEORIDER)
  • National Science Foundation (BCS-1523264)
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Forensic and Genetic Research
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic diversity and population structure
  • Social Sciences › Arts and Humanities › History › Eurasian Exchange Networks
Keywords
ancient population genetics, Xiongnu, Mongols, nomadic pastoralism, Mongolian population, population mixing, Bronze Age, Mongolian steppe
MeSH terms
Archaeology, Europe, Female, Gene Frequency, Gene Pool, Genetics, Population, Geography, Haplotypes, Humans, Male, Mongolia, Time Factors, Genome, Human, Genetic Heterogeneity, Principal Component Analysis, History, Ancient, Grassland

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

A language model drew the claims from the paper on 2026-11-12, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).

A second model labelled each claim on 2026-11-12 (model jev-1.13-20260917, definition revision gold-guideline-v0.2).

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

What counts as a claim ›How we measure confidence ›How we check this ›

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