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. 2023 Apr 14;9(15):eadf3904.
doi: 10.1126/sciadv.adf3904. Epub 2023 Apr 14.

Genetic population structure of the Xiongnu Empire at imperial and local scales

Juhyeon Lee  1 , Bryan K Miller  2   3   4 , Jamsranjav Bayarsaikhan  2   5 , Erik Johannesson  6 , Alicia Ventresca Miller  2   3   7 , Christina Warinner  8   9 , Choongwon Jeong  1
Affiliations

Genetic population structure of the Xiongnu Empire at imperial and local scales

Juhyeon Lee et al. Sci Adv. .

Abstract

The Xiongnu established the first nomadic imperial power, controlling the Eastern Eurasian steppe from ca. 200 BCE to 100 CE. Recent archaeogenetic studies identified extreme levels of genetic diversity across the empire, corroborating historical records of the Xiongnu Empire being multiethnic. However, it has remained unknown how this diversity was structured at the local community level or by sociopolitical status. To address this, we investigated aristocratic and local elite cemeteries at the western frontier of the empire. Analyzing genome-wide data from 18 individuals, we show that genetic diversity within these communities was comparable to the empire as a whole, and that high diversity was also observed within extended families. Genetic heterogeneity was highest among the lowest-status individuals, implying diverse origins, while higher-status individuals harbored less genetic diversity, suggesting that elite status and power was concentrated within specific subsets of the broader Xiongnu population.

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Figures

Fig. 1.
Fig. 1.. Map of Xiongnu sites in this study, and burial plans of the TAK and SBB cemeteries.
(A) Geographic locations of the sites analyzed in the study are presented with the cultural affiliation and the time period. Newly sequenced individuals were excavated from the aristocratic elite cemetery of Takhiltyn Khotgor (TAK) (yellow square) and the local elite cemetery of Shombuuzyn Belchir (SBB) (red circle) in western Mongolia. Previously reported Xiongnu sites with five or more sequenced individuals are also labeled in the figure: Salkhityn Am (SKT), Uguumur Uul (UGU), and the Il’movaya Pad (IMA) (14). Other Xiongnu sites are indicated with white diamonds (early Xiongnu) or black diamonds (late Xiongnu). Sites of the preceding Early Iron Age (EIA) associated with the Sagly/Uyuk (pink triangle) and Slab Grave (green down-pointing triangle) archaeological cultures are also shown. (B) Plan detail of the TAK cemetery indicating the square tomb complexes and associated graves (yellow). See fig. S1 for a full cemetery plan. (C) Plan of the SBB cemetery indicating stone circle and stone pile graves. Excavation focused on a dense grave cluster (pink) and a selection of other representative graves (red). See fig. S2 for a full cemetery plan. For both sites, tomb or grave numbers are indicated in bold red; each analyzed individual is numbered in black.
Fig. 2.
Fig. 2.. qpAdm modeling of TAK and SBB individuals.
The genetic profiles of ancient individuals are modeled as the mixture of two or three populations. The ancestry proportion of each source population is represented by the size of the box on the x axis. Horizontal bars represent ±1 SE estimated by qpAdm using 5-cM block jackknifing. Detailed results are presented in data file S2. (A) Ulaanzuukh and SlabGrave individuals are modeled as the mixture between Ancient Northeast Asian (ANA), represented by eastMongolia_preBA in this study, and Khovsgol_LBA. We modeled Takhiltyn Khotgor (TAK) (B) and Shombuuzyn Belchir (SBB) (C) individuals as the mixture of the preceding groups from Mongolia and the surrounding regions: SlabGrave1, Chandman_IA, Han_2000BP, Gonur1_BA, earlyXiongnu_west, Khovsgol_LBA, and UKY. We present two models for TAK008 and TAK009.
Fig. 3.
Fig. 3.. Genetic diversity of the Xiongnu.
(A) Principal components analysis (PCA) of Takhiltyn Khotgor (TAK) (yellow squares) and Shombuuzyn Belchir (SBB) (red circles) individuals. Other Xiongnu individuals are shown as hollow diamonds (early Xiongnu) and black diamonds (late Xiongnu). Ancient individuals were projected on the PCs calculated with 2077 present-day Eurasian individuals (gray). Inset shows PC1 on the x-axis and PC3 on the y axis. PC3 explains the 0.33% of the total variance. The x axis and y axis ticks have the same values across all panels, except the inset of (A) where the y axis ranges from −0.04 to 0.06. (B) PCA of Gol Mod 2 (azure squares) and other Xiongnu sites with five or more genetically analyzed individuals: Salkhityn Am (SKT) (purple squares), Uguumur Uul (blue circles), and Il’movaya Pad (IMA) (green circles). (C) PCA of the preceding Early Iron Age (EIA) Sagly/Uyuk (pink triangles) and Slab Grave (green down-pointing triangles) individuals. (D) PCA of TAK individuals, with color indicating tomb complex and size reflecting grave type. (E) PCA of SBB individuals, with color indicating cluster membership and size reflecting grave type. SBB005 is a second-degree genetic relative of SBB001 and SBB007.
Fig. 4.
Fig. 4.. Genetic diversity of the Xiongnu at local and imperial scales.
Left presents PC1 coordinates for individuals at Takhiltyn Khotgor (TAK), Shombuuzyn Belchir (SBB), and three previously reported Xiongnu sites with five or more sequenced individuals (14). For comparison, we also display PC1 coordinates for Xiongnu, Chandman_IA, and SlabGrave individuals as three independent clusters. The color of the boxes indicates the time period: Early Iron, pink; early Xiongnu, dark gray; late Xiongnu, light gray; and entire Xiongnu, light blue. Newly reported sites (TAK and SBB) are marked by a surrounding box. The PC1 distribution of Xiongnu individuals as a whole exceeds that of the prior Early Iron Age (EIA). With the exception of Il’movaya Pad (IMA), the PC1 distribution of individuals within each Xiongnu site is broader than that of either Chandman_IA or SlabGrave, and the overall PC1 distribution of individuals at Xiongnu sites is generally high. This indicates that the high genetic diversity observed among the Xiongnu as a whole is also reflected within Xiongnu communities. On the right, we compare PC1 coordinates of TAK/SBB individuals according to their social status (high versus low). The color of the boxes indicates social status: green, high; yellow, low; and it matches the color of the dots on the left. The overall PC1 distribution of the high-class individuals is more restricted than that of the lower class individuals.
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