‘Following the Xiongnu and Xianbei, the Rouran Khaganate (Rouran) was the third great nomadic tribe on the Mongolian Steppe. However, few human remains from this tribe are available for archaeologists and geneticists to study, as traces of the tombs of these nomadic peo- ple have rarely been found.
In 2014, the IA-M1 remains (TL1) at the Khermen Tal site from the Rouran period were found by a Sino-Mongolian joint archaeological team in Mongolia, providing precious material for research into the genetic imprint of the Rouran.
Materials and methods: The mtDNA hypervariable sequence I (HVS-I) and Y-chromosome SNPs were analyzed, and capture of the paternal non-recombining region of the Y chromosome (NRY) and whole-genome shotgun sequencing of TL1 were performed. The materials from three sites representing the three ancient nationalities (Donghu, Xianbei, and Shiwei) were selected for com- parison with the TL1 individual.
The mitochondrial haplotype of the TL1 individual was D4b1a2a1. The Y-chromosome haplotype was C2b1a1b/F3830 (ISOGG 2015), which was the same as that of the other two ancient male nomadic samples (ZHS5 and GG3) related to the Xianbei and Shiwei, which were also detected as F3889; this haplotype was reported to be downstream of F3830 by Wei et al. (2017).
We conclude that F3889 downstream of F3830 is an important paternal lineage of the ancient Donghu nomads. The Donghu-Xianbei branch is expected to have made an important paternal genetic contribution to Rouran. This component of gene flow ultimately entered the gene pool of modern Mongolic- and Manchu-speaking populations.
Since the Xiongnu and Han Dynasty successfully established unified political power on the Eurasian Steppes and Central Plains of China at the end of the third century BC, conflicts, exchanges and interactions between these two great civilizations (i.e., nomadic and farming civilizations) occurred over more than 2000 years of Chinese history.
Another nomadic tribe, the Donghu, lived in the east of Xiongnu. This tribe existed for 1300 years, from the early Shang Dynasty (1600 BC) to the Western Han Dynasty (206 BC) (Zhang et al., 2014). After Donghu was destroyed by Xiongnu, the remaining Donghu population evolved into two lineages (Wuhuan and Xianbei) (Lin, 1989).
The Xianbei lineage rapidly grew strong since the Xiongnu declined and moved westward. This lineage occupied the Mongolian Steppe, annexed the remaining tribes that once surrendered to Xiongnu, and eventually established the second largest nomadic Empire (Jing, 2013).
Following the Xiongnu and Xianbei, the Rouran Khaganate (Rouran) was the third great nomadic tribe in the Mongolian grasslands from the late fourth century to the mid-sixth century (Barbara, 2008). The Rouran controlled the area of Mongolia from the Manchurian bor- der to Turpan, perhaps even to the east coast of Lake Balkhash, ranging from the Orkhon River to China.
The core area was mainly located in the territory of modern Mongolia. The territory sometimes expanded west to the ancient Western Regions (now part of Central Asia and Xinjiang, China) and south to the northern part of China's Inner Mongolia Autonomous Region. Rouran set up a powerful nomadic empire and controlled the Silk Road, which ran through Yiwu, the Tianshan Mountains, Qiuci, and Pamirsfor ~150 years from the late fourth century to the mid-sixth century (Jing, 2013).
Shiwei was another ancient nomadic group that might originated from Xianbei, and this population mainly inhabited the Greater Khingan Mountains, dating from the fifth century (J, 1985). During the 10th century, the Shiwei population began to migrate westward, finally reaching the region currently known as Mongolia (J, 1985; Zhang et al., 2018).
However, the origins of nomadic tribes such as the Rouran that emerged after the Xianbei are puzzling because historical records concerning these tribes are rare.The customs and lifestyles differed between the nomadic population and the farming population.
Most of the nomads lived in grasslands and did not settle, and there were no special signs indicating their tombs; thus, few remains are available for archaeologists and geneticists to study. In 2014, a Sino-Mongolian joint archaeological team excavated a small earthen pit tomb at the Khermen Tal site in Ogiinuur Sum in Arkhangai Province, Mongolia (Figure 1a) (Chen et al., 2015a).
The funerary objects unearthed from this tomb are abundant and include pottery, gold headdresses, arched bronze pendants, bronze bracelets, bronze rings, bronze makeup appliances, leather bags, silk fabric, iron belts, iron arrowheads, iron swords and bows made of bone.
These objects are relatively rare in tombs characterized by nomadic cultures in Mongolia. The pottery in this tomb exhibits special shape characteristics and displays iconic artifacts of the Xianbei culture (Zheng, 2009). The arched bronze pendant on the neck of the tomb's owner was similar to the pendant found at the Chenwugou Xianbei site in Huade county, Inner Mongolia Autonomous Region (Hu et al., 2014).
The 14C dating of the wood on the coffin board was confirmed both at Peking University, China (cal. AD 590-655) and the University of Cologne, Germany (cal. AD 335-535) (Chen et al., 2015a). The absolute chronology of the tomb ranges from the fourth to sixth centuries AD, which is a time period coinciding with the Rouran Khan Kingdom on the Mongolian Plateau.
The article by Chen et al. (2015a) confirmed that this site was a Rouran site, based on the unearthed funerary objects, the absolute chronology, and the unique cultural attributes and connotations of this tomb. The tomb was the first of this period to be found in Mongolia and has provided us with precious material for research into the genetic imprint of the Rouran.
The mDNA hypervariable sequence | (HVS-I) and Y-chromosome SNPs were analyzed, and capture of the paternal non-recombining region of the Y chromosome (NRY) and whole-genome shotgun sequencing were performed on the IA-M1 remains (TL1) from the Khermen Tal site.
To study the genetic imprint of the Rouran Khaga-nate and the paternal genetic characteristics of the Donghu-Xianbei lin-eage, eight additional samples from three sites (the Jinggouzi, Zaan khoshuu, and Gangga sites) that are representative of three ancient nomadic tribes (Donghu, Xianbei, and Shiwei) were selected for comparison with the TL1 individual.
Phylogenetic analysis of the Y chromosome
The phylogenetic tree constructed using the NRY capture data from the three ancient samples and 25 modern reference samples belonging to haplogroup C2 (proto C3) (Supporting Information S5) (Lippold et al.,
2014; Wei et al., 2017) showed that the ancient individuals were located together on a branch (F3889) with modern Mongolian-Buryats, Xibo, Yugur, Hezhen, Kazakh, and Han individuals.
Analysis of the BSP for the NRY data showed that Hg C2 (proto C3) experienced a bottleneck ~2000 years ago and that the effective male population size increased two times (~500-year-old and ~1500-year-old) over the subsequent 2000 years.
Genetic imprint of the Rouran Khaganate
In our research, the mitochondrial haplotype of the TL1 individual was D4b1a2a1, which is downstream of haplogroup D4, the most frequently occurring mDNA haplogroup among modern populations of northern East Asia, including the Japanese, Okinawans, Koreans, and Mongolic- or Tungusic-speaking populations of northern China (Der-enko et al., 2012; Kong et al., 2003; Lee et al., 2006; Maruyama, Mina-guchi, & Saitou, 2003; Umetsu et al., 2005; Zheng et al., 2011).
Previous studies have also shown that haplogroup D4 was the most common haplogroup among the ancient populations of northern China, such as those associated with the Qilangshan, Dondajing, and Lama-dong sites (Wang et al., 2007; Yu, Xie, Zhang, Zhou, & Zhu, 2006; Yu, Zhao, & Zhou, 2014).
The research of Derenko et al. (2010) implied that D4b1a2a1, downstream of haplogroup D4b1a2, originated in southern Siberia, and the age of D4b1a2a1 was estimated at ~11kya (Derenko et al., 2010). From the phylogenetic tree of the mitochondrial genomes, we found that haplogroup D4b1a2a1 of the TL1 individual was mainly distributed in Russia and Inner Mongolia, China.
Combined with the results of previous research, we speculate that mDNA hap-logroup D4b1a2a1 most likely originated from the Mongolian grass-lands. In addition, PCA based on nuclear data showed that the ancient TL1 individual shared high similarity with contemporary Yakut and Mongola populations.
We also observed that TL1 exhibited a very similar composition to the Yakut, Oroqen, Daur, Hezhen, and Mongola populations according to admixture analysis. Moreover, Treemix and f3 test analyses revealed that TL1 displayed a close genetic relationship to the Oroqen and Daur populations. All of these results indicated that the TL1 individual made an important genetic contribution to modern Mongolic- and Manchu-speaking peoples.
The Y-chromosome haplotype of the TL1 individual was F3889, positioned downstream of C2b1a1b/F3830 (ISOGG 2015), which accounts for a very low proportion of haplogroup C2b1a/F3918. The Y-chromosome haplotypes of ZHS5 (from the Zaan khoshuu site) and GG3 (from the Gangga site) also belonged to F3889. The ZHS5 sample, from the Zaan khoshuu site, came from the Xianbei period.
The Gangga people belonged to the Shiwei culture, which might also have originated from the Xianbei population. All of these people were related to the ancient Donghu-Xianbei branch. These results suggested that TL1 likely presents a close paternal relationship to the Donghu people and may have even descended from a branch of the ancient Donghu-Xianbei people, based on the conclusion that haplogroup C2b1a/F3918 can be considered the paternal branch of the ancient Donghu people (Zhang et al., 2018).
The Y-chromosome phylogenetic tree showed that TL1 shared a branch with modern Mongolian-Buryats, Hezhen, Xibo, Yugur, and Kazakh, suggesting that the TL1 individual from the Rouran period should also generally present close paternal genetic relationships with modern Mongolic-and Manchu-speaking peoples.
In general, the Rouran Khaganate originated from an alliance of the ancient Eurasian steppe nomads, which disintegrated and disappeared with the progress of history. This group was complex, and its origin cannot be explained based only on one individual. However, we can trace the genetic imprint of the Rouran people through genome analysis of the TL1 individual.
On the basis of the comparison with other ancient nomadic people (Donghu, Xianbei, and Shiwei) and data on modern individuals from published articles (Lippold et al., 2014; Wei et al., 2017) (Supporting Information S5), we found that they all share the same haplotype implying shared paternal ancestry between the Donghu, Xianbei and Rouran populations.
Furthermore, this gene flow (mainly haplogroup C2b1a/F3918) did not stop with the disappearance of the Rouran, and a portion was instead passed on in other groups, such as the ancient Shiwei people (later than Rouran), eventually reaching the gene pool of modern Mongolic- and Manchu-speaking populations (Mongolian-Buryats, Hezhen, Xibo, et al).
Important paternal lineage in Donghu nomads:
F3889 Hg C3* is widely distributed in Eurasia, as an important paternal lineage with a typical paternal haplogroup of Mongolic-speaking populations, and shows a moderate distribution among other Tungusic peoples (Malyarchuk et al., 2010).
This group is not monophyletic and has several sub-haplogroups, among which the C3*-Star cluster (C2b1c/ F1918, ISOGG 2015) presents an extremely high frequency in the Mongolian tribes and Turkic peoples and is thought to be the paternal haplotype of Gen Gi Khan or his male relatives (Zerjal et al., 2003).
The C3*-DYS448del (C2b1a1b/F1756, ISOGG 2015) cluster is widely distributed at a low frequency in almost all Mongolic- and Turkic-speaking populations. Wei et al. (2017) proposed that the C3*-DYS448del cluster might be a candidate for the paternal lineages of the ancient Donghu, Xianbei, and Shiwei tribes (Wei et al., 2017).
However, these authors lacked data on ancient DNA to verify this hypothesis. Through the NRY capture analysis conducted in the present study, the TL1 individual and other two ancient samples [ZHS5 (~1400 years old, belonging to Xianbei) and GG3 (~1100 years old, belonging to Shiwei)] were allocated to haplotype F3889, which was the lower part of C2b1a1b/ F1756 (proto C3*-DYS448del).
Moreover, we found that C3*-Star separated early from C3*-DYS448del in the Y-chromosome phylogenetic tree, whereas the divergence time between C3*-DYS448del and its most closely related lineage was ~13 kya. The expansion time for branch F3889 (~3.3 kya), downstream of C3*-DYS448del, was much earlier than that of the C3*-Star cluster (~2.8 kya). This period corresponded only to the reign of the ancient Donghu nomads (from the early Shang Dynasty to the Western Han Dynasty) in northeast China.
Our results supported the inference drawn by Wei et al. (2017) and further indicated that the F3889 branch, downstream of C2b1a1b/ F1756 (proto C3*-DYS448del), was an important paternal lineage in Donghu-Xianbei nomads before the expansion of the Mongols.
Source: The genome of an ancient Rouran individual reveals an important paternal lineage in the Donghu population" - Jiawei Li, Ye Zhang, Yongbin Zhao, Yongzhi Chen, A. Ochir, Sarenbilige, Hong Zhu, Hui Zhou
Authors: Jiawei Li, Ye Zhang, Yongbin Zhao, Yongzhi Chen, A. Ochir, Sarenbilige, Hong Zhu, Hui Zhou
Institutions: Ancient DNA Laboratory, Research Center for Chinese Frontier Archaeology, Jilin University, Changchun, People’s Republic of China, College of Life Science, Jilin University, Changchun, People’s Republic of China, Life Science College, Jilin Normal University, Siping 136000, People’s Republic of China, Director, Inner Mongolian Museum, Hohhot, People’s Republic of China, Coordinator, International Institute for Study of Nomadic Civilization, Ulaanbaatar, Mongolia, Editorial department, Cultural Relics and Archaeological Institute of Inner Mongolia, Hohhot, People’s Republic of China, Correspondence Hui Zhou, College of Life Science, Jilin University, Changchun, People’s Republic of China.
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