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The deep population history of East Asia remains poorly understood due to a lack of ancient DNA data and sparse sampling of present-day people. We report genome-wide data from 166 East Asians dating to 6000 BCE – 1000 CE and 46 present-day groups. Hunter-gatherers from Japan, the Amur River Basin, and people of Neolithic and Iron Age Taiwan and the Tibetan plateau are linked by a deeply-splitting lineage likely reflecting a Late Pleistocene coastal migration. We follow Holocene expansions from four regions. First, hunter-gatherers of Mongolia and the Amur River Basin have ancestry shared by Mongolic and Tungusic language speakers but do not carry West Liao River farmer ancestry contradicting theories that their expansion spread these proto-languages. Second, Yellow River Basin farmers at ~3000 BCE likely spread Sino-Tibetan languages as their ancestry dispersed both to Tibet where it forms up ~84% to some groups and to the Central Plain where it contributed ~59-84% to Han Chinese. Third, people from Taiwan ~1300 BCE to 800 CE derived ~75% ancestry from a lineage also common in modern Austronesian, Tai-Kadai and Austroasiatic speakers likely deriving from Yangtze River Valley farmers; ancient Taiwan people also derived ~25% ancestry from a northern lineage related to but different from Yellow River farmers implying an additional north-to-south expansion. Fourth, Yamnaya Steppe pastoralist ancestry arrived in western Mongolia after ~3000 BCE but was displaced by previously established lineages even while it persisted in western China as expected if it spread the ancestor of Tocharian Indo-European languages. Two later gene flows affected western Mongolia: after ~2000 BCE migrants with Yamnaya and European farmer ancestry, and episodic impacts of later groups with ancestry from Turan.

01. Genomic Insights into the Formation of Human Populations in East Asia

Nature, 22 February 2021

https://doi. org/10.1038/s41586-021-03336-2 (2021).

Chuan-Chao Wang, Hui-Yuan Yeh, Alexander N. Popov, Hu-Qin Zhang, Hirofumi Matsumura, Kendra Sirak, Olivia Cheronet, Alexey Kovalev, Nadin Rohland, Alexander M. Kim, Swapan Mallick, Rebecca Bernardos, Dashtseveg Tumen, Jing Zhao, Yi-Chang Liu, Jiun-Yu Liu, Matthew Mah, Ke Wang, Zhao Zhang, Nicole Adamski, Nasreen Broomandkhoshbacht, Kimberly Callan, Francesca Candilio, Kellie Sara Duffett Carlson, Brendan J. Culleton, Laurie Eccles, Suzanne Freilich, Denise Keating, Ann Marie Lawson, Kirsten Mandl, Megan Michel, Jonas Oppenheimer, Kadir Toykan Özdoğan, Kristin Stewardson, Shaoqing Wen, Shi Yan, Fatma Zalzala, Richard Chuang, Ching-Jung Huang, Hana Looh, Chung-Ching Shiung, Yuri G. Nikitin, Andrei V. Tabarev, Alexey A. Tishkin, Song Lin, Zhou-Yong Sun, Xiao-Ming Wu, Tie-Lin Yang, Xi Hu, Liang Chen, Hua Du, Jamsranjav Bayarsaikhan, Enkhbayar Mijiddorj, Diimaajav Erdenebaatar, Tumur-Ochir Iderkhangai, Erdene Myagmar, Hideaki Kanzawa-Kiriyama, Masato Nishino, Ken-ichi Shinoda, Olga A. Shubina, Jianxin Guo, Wangwei Cai, Qiongying Deng, Longli Kang, Dawei Li, Dongna Li, Rong Lin, Nini, Rukesh Shrestha, Ling-Xiang Wang, Lanhai Wei, Guangmao Xie, Hongbing Yao, Manfei Zhang, Guanglin He, Xiaomin Yang, Rong Hu, Martine Robbeets, Stephan Schiffels, Douglas J. Kennett, Li Jin , Hui Li, Johannes Krause, Ron Pinhasi & David Reich




02. Ancient DNA and multimethod dating confirm the late arrival of anatomically modern humans in southern China

PNAS, 23 February 2021

2021: 118 (8) e2019158118;

Xue-feng Sun, Shao-qing Wen, Cheng-qiu Lu, Bo-yan Zhou, Darren Curnoe, Hua-yu Lu, Hong-chun Li, Wei Wang, Hai Cheng, Shuang-wen Yi, Xin Jia, Pan-xin Du, Xing-hua Xu, Yi-ming Lu, Ying Lu, Hong-xiang Zheng, Hong Zhang, Chang Sun, Lan-hai Wei, Fei Han, Juan Huang, R. Lawrence Edwards, Li Jin, and Hui Li


The expansion of anatomically modern humans (AMHs) from Africa around 65,000 to 45,000 y ago (ca. 65 to 45 ka) led to the establishment of present-day non-African populations. Some paleoanthropologists have argued that fossil discoveries from Huanglong, Zhiren, Luna, and Fuyan caves in southern China indicate one or more prior dispersals, perhaps as early as ca. 120 ka. We investigated the age of the human remains from three of these localities and two additional early AMH sites (Yangjiapo and Sanyou caves, Hubei) by combining ancient DNA (aDNA) analysis with a multimethod geological dating strategy. Although U–Th dating of capping flowstones suggested they lie within the range ca. 168 to 70 ka, analyses of aDNA and direct AMS 14C dating on human teeth from Fuyan and Yangjiapo caves showed they derive from the Holocene. OSL dating of sediments and AMS 14C analysis of mammal teeth and charcoal also demonstrated major discrepancies from the flowstone ages; the difference between them being an order of magnitude or more at most of these localities. Our work highlights the surprisingly complex depositional history recorded at these subtropical caves which involved one or more episodes of erosion and redeposition or intrusion as recently as the late Holocene. In light of our findings, the first appearance datum for AMHs in southern China should probably lie within the timeframe set by molecular data of ca. 50 to 45 ka.

03. Multidisciplinary evidence supported the hypothesis of north China origin of Sino-Tibetan



Chinese Social Sciences Today (中国社会科学报 ), 11 May 2021

Jian-xin GuoChuan-chao Wang


The origin and dispersal of the Sino-Tibetan has been one of the most concerned question in historical comparative linguistics. It noted that the comprehensive analysis of archaeological defined cultures and genetically attested population relationships showed accurate illustration in Sino-Tibetan research, providing multidisciplinary evidence for the origin and dispersal from northern East Asia. Focus on “Farming-language/ethnic dispersal Hypothesis”, linguistic and archaeological studies outlined the migration route of proto-Sino-Tibetan originated from the North China and then dispersed to middle and upper Yellow River area, Southwest China and the Southeast Asia, and supported the hypothesis of “Tibetan-Yi Corridor” in Southwest China since Neolithic Period. By capitalizing on time transects of ancient DNA research in East Asia, we observed the strong genetic affinity between millet-farming-related ancients and modern Sino-Tibetan, which is consistent with the cultural links reflected in linguistic and archaeological studies. Furthermore, from genome-wide ancient DNA perspective, we observed large-scale genetic contribution in modern Sino-Tibetan associated Yangshao and Majiayao Culture ancients, suggesting continuous genetic flow from northern China millet farmers. Therefore, the population migrations in Tibetan-Yi Corridor in southwest East Asia supported by paleogenomicsevidences are coincident with the linguistically and archaeologically attested proto-Sino-Tibetan expansions associated with “Farming-language/ethnic dispersal Hypothesis”.

The work was funded by the European Research Council (ERC), ERC-2019-AdG-883700-TRAM

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04. Origin of ethnic groups, linguistic families, and civilizations in China

viewed from the Y chromosome

Molecular Genetics and Genomics , 26 May 2021

Xueer YuHui Li


East Asia, geographically extending to the Pamir Plateau in the west, to the Himalayan Mountains in the southwest, to Lake Baikal in the north and to the South China Sea in the south, harbors a variety of people, cultures, and languages. To reconstruct the natural history of East Asians is a mission of multiple disciplines, including genetics, archaeology, linguistics, and ethnology. Geneticists confirm the recent African origin of modern East Asians. Anatomically modern humans arose in Africa and immigrated into East Asia via a southern route approximately 50,000 years ago. Following the end of the Last Glacial Maximum approximately 12,000 years ago, rice and millet were domesticated in the south and north of East Asia, respectively, which allowed human populations to expand and linguistic families and ethnic groups to develop. These Neolithic populations produced a strong relation between the present genetic structures and linguistic families. The expansion of the Hongshan people from northeastern China relocated most of the ethnic populations on a large scale approximately 5300 years ago. Most of the ethnic groups migrated to remote regions, producing genetic structure differences between the edge and center of East Asia. In central China, pronounced population admixture occurred and accelerated over time, which subsequently formed the Han Chinese population and eventually the Chinese civilization. Population migration between the north and the south throughout history has left a smooth gradient in north–south changes in genetic structure. Observation of the process of shaping the genetic structure of East Asians may help in understanding the global natural history of modern humans.

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05. Tracing the Case Forming in Sinitic Languages and Varieties in the Gansu-Qinghai Linguistic Area


Linguistic Sciences (语言科学) , May 2021


Dan Xu


By tracing and comparing the corpus from the 1960s to the present, we have found that the case marking in Sinitic languages and varieties has been formed gradually. Amdo Tibetan, Mongolic, and Turkic languages have influenced the Sinitic languages at different periods and locations. The case system we can observe today constitutes a competition result from different languages. The accusative case marking is overwhelmingly attested in this area. It is clear that the type of accusative case marking used in Mongolic and Turkic languages prevails over that of ergative case marking seen in Tibetan languages. At the beginning time where Sinitic languages were changing the word order from VO to OV, preverbal direct objects, as well as indirect objects, were not marked by any case. The Hezheng dialect was the first to use an accusative/dative marker with a phonetic form [a] in the corpus from the 1960s. It seems that the postposition [xa] ‘under’ in Sinitic languages could not be the source of this marker: it is a type of borrowing based on pattern models from other languages, while the phonetic form comes from the topic marker [a/xa] in Sinitic languages. In possession and experiencer sentences, the dative marker use can be found in published data after 80s. Our study shows that this dative marker has been formed later than the first type. As for the pronominal marking, the first and second person pronouns have performed earlier the case marking than the third person pronoun; moreover, the first and second person pronouns have experienced a coalescence with the case marker.

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06. The Source of polokai ‘knee’ in Chinese: Deep Contact between the Chinese and Non-Chinese Languages

(汉语“波罗盖(膝)”的来源 ——兼谈汉语与非汉语的深层接触)

Minority Languages of China (民族语文) , 03, 2021


Dan Xu


This paper proposes that the word polokai ‘knee’, widely distributed in the dialects of northern China, did not have its source in the Chinese language itself, but come from substrate languages spoken by northern populations. It was probably transmitted through the Jurchenized Tungusic people. Historical documents show that during the formation of the Jurchen and Manchu, diverse tribes had been integrated into them. The linguistic data shows that the p-l-k (polokai ‘knee’) pattern in the northern Chinese dialects is likely to be related to the b-l-g ‘knee’ pattern in Tungusic languages. The morphological means observed in Tungusic languages may have a deep connection with that found in languages of nomadic, fishing-hunting populations in the North. This word is probably linked to a cognate root bul- / pal- shared between these ancient languages.

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07. Exploring the origin and diversification of the Xiongnu from a multidisciplinary perspective



Chinese Social Sciences Today (中国社会科学报 ), 13 Septembre 2021

Na SunChuan-chao Wang

The work was funded by the European Research Council (ERC), ERC-2019-AdG-883700-TRAM

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08. Genome‐wide autosomal, mtDNA, and Y chromosome analysis of King Bela III of the Hungarian Arpad dynasty

Scientific Reports, (2021) 11: 19210

Chuan-Chao Wang, Cosimo Posth, Anja Furtwängler, Katalin Sümegi, Zsolt Bánfai, Miklós Kásler, Johannes Krause, & Béla Melegh





The ancient Hungarians, “Madzsars”, established their control of the Carpathian Basin in the late ninth century and founded the Hungarian Kingdom around 1000AD. The origin of the Magyars as a tribal federation has been much debated in the past. From the time of the conquest to the early fourteenth century they were ruled by descendants of the Arpad family. In order to learn more about the genetic origin of this family, we here analyzed the genome of Bela III one of the most prominent members of the early Hungarian dynasty that ruled the Hungarian Kingdom from 1172 to 1196. The Y‐Chromosome of Bela III belongs to haplogroup R1a‐Z2123 that is today found in highest frequency in Central Asia, supporting a Central Asian origin for the ruling lineage of the Hungarian kingdom. The autosomal DNA profile of Bela III, however, falls within the genetic variation of present‐day east European populations. This is further supported through his mtDNA genome that belongs to haplogroup H, the most common European maternal lineage, but also found in Central Asia. However, we didn’t find an exact haplotype match for Bela III. The typical autosomal and maternal Central Eastern European ancestry among Bela III autosomes might be best explained by consecutive intermarriage with local European ruling families.

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09. Genomic Insight Into the Population Structure and Admixture History of Tai-Kadai-Speaking Sui People in Southwest China

Frontiers in Genetics, 20 Septembre 2021

doi: 10.3389/fgene.2021.735084

Xiaoyun Bin, Rui Wang, Youyi Huang, Rongyao Wei, Kongyang Zhu, Xiaomin Yang, Hao Ma, Guanglin He, Jianxin Guo, Jing Zhao,
Meiqing Yang, Jing Chen, Xianpeng Zhang, Le Tao, Yilan Liu, Xiufeng Huang and
Chuan-Chao Wang





Sui people, which belong to the Tai-Kadai-speaking family, remain poorly characterized due to a lack of genome-wide data. To infer the fine-scale population genetic structure and putative genetic sources of the Sui people, we genotyped 498,655 genome-wide single-nucleotide polymorphisms (SNPs) using SNP arrays in 68 Sui individuals from seven indigenous populations in Guizhou province and Guangxi Zhuang Autonomous Region in Southwest China and co-analyzed with available East Asians via a series of population genetic methods including principal component analysis (PCA), ADMIXTURE, pairwise Fst genetic distance, f-statistics, qpWave, and qpAdm. Our results revealed that Guangxi and Guizhou Sui people showed a strong genetic affinity with populations from southern China and Southeast Asia, especially Tai-Kadai- and Hmong-Mien- speaking populations as well as ancient Iron Age Taiwan Hanben, Gongguan individuals supporting the hypothesis that Sui people came from southern China originally. The indigenous Tai-Kadai-related ancestry (represented by Li), Northern East Asian-related ancestry, and Hmong-Mien-related lineage contributed to the formation processes of the Sui people. We identified the genetic substructure within Sui groups: Guizhou Sui people were relatively homogeneous and possessed similar genetic profiles with neighboring Tai-Kadai-related populations, such as Maonan. While Sui people in Yizhou and Huanjiang of Guangxi might receive unique, additional gene flow from Hmong- Mien-speaking populations and Northern East Asians, respectively, after the divergence within other Sui populations. Sui people could be modeled as the admixture of ancient Yellow River Basin farmer-related ancestry (36.2–54.7%) and ancient coastal Southeast Asian-related ancestry (45.3–63.8%). We also identified the potential positive selection signals related to the disease susceptibility in Sui people via integrated haplotype score (iHS) and number of segregating sites by length (nSL) scores. These genomic findings provided new insights into the demographic history of Tai-Kadai-speaking Sui people and their interaction with neighboring populations in Southern China.

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10. On the disjunctive constructions and related constructions in Zhoutun

Lingua, 24 Septembre 2021

Chenlei Zhou


This paper focuses on two disjunctive constructions in Zhoutun, namely, the puʂi-disjunctive construction “[[(puʂi) C1…, puʂi Cn]←→[Alternation: C1- Cn]]” and the -disjunctive construction “[[X  Y]←→[Alternation: X-Y]]”. The prepositional, standard disjunctive coordinator puʂi links declarative clausal coordinands, and the postpositional, intterogative disjunctive coordinator  links clauses and NPs. The constructionalization process of the two constructions is discussed in detail. The puʂi-disjunctive construction underwent the process of “C1, tʂɤmɤ puʂi ʂi, C2 > C1, pu ʂi ʂi, C2 > C1, puʂi C2 > (puʂi) C1…, puʂi Cn”. The -disjunctive construction originated from the -question construction in which  is the question marker. Another two related constructions containing puʂi, the puʂi-imperative construction “[[puʂi Cimp]←→[Imperative with softer tone]]” and the puʂi-exceptive construction “[[NP puʂi (ʂiNEG]←→[Only NP]]” are also discussed in terms of their use and constructionalization process.


11. Genomic Insight Into the Population Admixture History of Tungusic-Speaking Manchu People in Northeast China

Frontiers in Genetics, 30 Septembre 2021


Xianpeng Zhang, Guanglin He, Wenhui Li, Yunfeng Wang, Xin Li, Ying Chen, Quanying Qu, Ying Wang, Huanjiu Xi, Chuan-Chao Wang and Youfeng Wen 





Manchu is the third-largest ethnic minority in China and has the largest population size among the Tungusic-speaking groups. However, the genetic origin and admixture history of the Manchu people are far from clear due to the sparse sampling and a limited number of markers genotyped. Here, we provided the first batch of genome-wide data of genotyping approximate 700,000 single-nucleotide polymorphisms (SNPs) in 93 Manchu individuals collected from northeast China. We merged the newly generated data with data of publicly available modern and ancient East Asians to comprehensively characterize the genetic diversity and fine-scale population structure, as well as explore the genetic origin and admixture history of northern Chinese Manchus. We applied both descriptive methods of ADMIXTURE, fineSTRUCTURE, FST, TreeMix, identity by decedent (IBD), principal component analysis (PCA), and qualitative f-statistics (f3 , f4, qpAdm, and qpWave). We found that Liaoning Manchus have a close genetic relationship and significant admixture signal with northern Han Chinese, which is in line with the cluster patterns in the haplotype-based results. Additionally, the qpAdm-based admixture models showed that modern Manchu people were formed as major ancestry related to Yellow River farmers and minor ancestry linked to ancient populations from Amur River Bain, or others. In summary, the northeastern Chinese Manchu people in Liaoning were an exception to the coherent genetic structure of Tungusic-speaking populations, probably due to the large-scale population migrations and genetic admixtures in the past few hundred years.

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12. Comparative Performance of the MGISEQ-2000 and Illumina X-Ten Sequencing Platforms for Paleogenomics

Frontiers in Genetics, 04 Octobre 2021


Kongyang Zhu, Panxin Du, Jianxue Xiong, Xiaoying Ren, Chang Sun, Yichen Tao, Yi Ding, Yiran Xu, Hailiang Meng, Chuan-Chao Wang and Shao-Qing Wen 





The MGISEQ-2000 sequencer is widely used in various omics studies, but the performance of this platform for paleogenomics has not been evaluated. We here compare the performance of MGISEQ-2000 with the Illumina X-Ten on ancient human DNA using four samples from 1750 BCE to 60 CE. We found there were only slight differences between the two platforms in most parameters (duplication rate, sequencing bias, θ, δS, and λ). MGISEQ-2000 performed well on endogenous rate and library complexity although X-Ten had a higher average base quality and lower error rate. Our results suggest that MGISEQ-2000 and X-Ten have comparable performance, and MGISEQ-2000 can be an alternative platform for paleogenomics sequencing.

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13. Inferring the population structure and admixture history of three Hmong-Mien-speaking Miao tribes from southwest China based on Genome-wide SNP genotyping

Annals of Human Biology, 12 November 2021

Ting Luo, Rui Wang, Chuan-Chao Wang 





Hmong-Mien speaking Miao, also called Hmong, is the sixthlargest ethnic group in mainland China. However, the fine-scale genetic profiles and population history of Miao populations in southwest China, especially in Guizhou province, remain uncharacterised due to a scarcity of samples of genome-wide data from different tribes. To further investigate the population substructure and admixture history of the Guizhou Miao minority. We collected 29 samples from three Miao tribes of Guizhou province in southwest China and genotyped about 700,000 genome-wide SNPs of each sample. We analysed newly generated data in together with published modern/ancient East Asian populations datasets via a series of population genetic methods, including principal component analysis (PCA), ADMIXTURE, Fst, TreeMix, f-statistics, qpWave, and qpAdm. PCA and ADMIXTURE results showed that the three studied Guizhou Miao groups consistently fell on the Hmong-Mien-related genetic cline and were relatively genetically homogeneous, displayingd a genetic affinity with neighbouring Tai-Kadai speaking populations such as Dong. These results were further confirmed by the observed genetic clade in Fst, TreeMix, outgroup-f3 -statistics, and f4-statistics. Furthermore, f4 -based allele sharing patterns illustrated that compared with Hunan Miao in central China, Guizhou Miao shared more alleles with Hmong-Mien-speaking Vietnam Hmong and Tai-Kadai-speaking CoLao, Dong, while exhibiting less northeast Asian-related ancestry. Admixture-f3 and f4 statistics revealed the North-South admixture pattern for the studied Guizhou Miao. A qpAdm-based two-way admixture model further revealed that the studied Guizhou Miao harboured 44%∼55.4% indigenous Austronesian-speaking Atayal-related ancestry and 44.6%∼56% Late Neolithic Yellow River farmer-related ancestry. The population structure within Hmong-Mien-related populations showed a geographic correlation. Hmong-Mien speaking Hunan Miao, Guizhou Miao, and Vietnam Hmong presented closer genetic relationships although they dwelt in different regions, suggesting the preservation of the original Hmong-related genetic diversity. The results based on genome-wide SNPs data generally matched the migration history for the Miao population. Our study contributes to a better knowledge of Miao populations and the population structure in southwest China.

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14. Genomic insights into Neolithic farming-related migrations in the junction of east and southeast Asia

American Journal of Biological Anthropology, 15 November 2021

Jianxin Guo, Weitao Wang, Kai Zhao, Guangxing Li, Guanglin He, Jing Zhao, Xiaomin Yang, Jinwen Chen, Kongyang Zhu, Rui Wang, Hao Ma, Bingying Xu, Chuan-Chao Wang 





Objectives: We aim to detect demographic history and early farming-related migration of the crossroad area in the junction of east Asia (EA) and southeast Asia (SEA).
Materials and methods: We collected and genotyped 87 individuals from 6 Tibeto-Burman and Austroasiatic-speaking populations including Bai, Pumi, Hani, Lahu, Wa, and Blang with nearly 700,000 genome-wide SNPs. We subsequently ana- lyzed genetic structure and admixture using our merged dataset including both ancient and modern eastern Eurasians with PCA, ADMIXTURE, Refine-IBD, statistics and qpAdm.
Results: We observed population substructure within the studied Tibeto-Burman populations. The northern Tibeto-Burman groups (Bai and Pumi) had a predominant genomic legacy associated to millet-farming from North China and also high frequen- cies of Y-chromosomal haplogroup O2a2b1-M134 (xM117) and its sub-clades. By contrast, southern Tibeto-Burman groups (Lahu and Hani) had more than 60% geno- mic legacy associated to rice-farming, which is prevalent in present-day Tai-Kadai, Austronesian and Austroasiatic speaking populations. We observed strong genetic affinities between Austroasiatic populations in Yunnan (Blang and Wa) and mainland southeast Asia.
Discussion: Our study revealed that both demic migrations and cultural interactions from north to south and east to west since the Late Neolithic have shaped the genetic structure of populations at the crossroads of EA and SEA. The dominant genomic legacy associated with millet-farming in northern Tibeto-Burman populations indicates large-scale Neolithic migrations from the Upper-Middle Yellow River Basin. The rice-farming expansion has deeply influenced the genetic profile of both southern Tibeto-Burman and Austroasiatic populations, suggesting migrations from east to west via both inland and coastal routes.

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15. Genomic insights into the genetic structure and natural selection of Mongolians

Frontiers in Genetics, November 2021


Xiaomin Yang, Sarengaowa Sarengaowa, Guanglin He, Jianxin Guo, Kongyang Zhu, Hao Ma, Jing Zhao, Meiqing Yang, Jing Chen, Xianpeng Zhang, Le Tao, Yilan Liu, Xiu-Fang Zhang and Chuan-Chao Wang 





Mongolians dwell at the Eastern Eurasian Steppe, where is the agriculture and pasture interlaced area, practice pastoral subsistence strategies for generations and have their own complex genetic formation history. There is evidence that the eastward expansion of Western Steppe herders transformed the lifestyle of post-Bronze Age Mongolia Plateau populations, and brought gene flow into the gene pool of Eastern Eurasians. Here, we reported genome-wide data for 42 individuals from the Inner Mongolia Autonomous Region of North China. We observed that our studied Mongolians were structured into three distinct genetic clusters possessing different genetic affinity with previous studied inner Mongolians and Mongols and various Eastern and Western Eurasian ancestry: two subgroups harbored dominant Eastern Eurasian ancestry from Neolithic millet farmers of Yellow River Basin; another subgroup derived Eastern Eurasian ancestry primarily from Neolithic hunter-gatherers of North Asia. Besides, three-way/four-way qpAdm admixture models revealed both north and southern Western Eurasian ancestry related to the Western Steppe herders and Iranian farmers contributed to the genetic materials into modern Mongolians. ALDER-based admixture coefficient and haplotype-based GLOBETROTTER demonstrated that the former western ancestry detected in modern Mongolian could be recently traced back to historic period in accordance with the historical record about the westward expansion of Mongol empire. Furthermore, the natural selection analysis of Mongolian showed that Major Histocompatibility Complex (MHC) region underwent significantly positive selective sweeps. The functional genes alcohol dehydrogenase (ADH) and lactase persistence (LCT) were not identified, while the higher/lower frequencies of derived mutations were strongly correlated with the genetic affinity to East Asian/Western Eurasian. Our attested complex population movement and admixture in the agriculture and pasture interlaced area played an important role in the formation of modern Mongolians.

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16. Genomic history and forensic characteristics of Sherpa highlanders on the Tibetan Plateau inferred from high-resolution InDel panel and genome-wide SNPs

Forensic Science International: Genetics, 18 November 2021

Mengge Wang, Weian Du, Renkuan Tang, Yan Liu, Xing Zou, Didi Yuan, Zheng Wang, Jing Liu, Jianxin Guo, Xiaomin Yang, Jing Chen, Meiqing Yang, Xianpeng Zhang,
Lan-Hai Wei, Haibing Yuan, Hui-Yuan Yeh, Chuan-Chao Wang, Chao Liu, Guanglin He 





Sherpa people, one of the high-altitude hypoxic adaptive populations, mainly reside in Nepal and the southern Tibet Autonomous Region. The genetic origin and detailed evolutionary profiles of Sherpas remain to be further explored and comprehensively characterized. Here we analyzed the newly-generated InDel genotype data from 628 Dingjie Sherpas by merging with 4222 worldwide InDel profiles and collected genome-wide SNP data (approximately 600 K SNPs) from 1612 individuals in 191 modern and ancient populations to explore and reconstruct the fine-scale genetic structure of Sherpas and their relationships with nearby modern and ancient East Asians based on the shared alleles and haplotypes. The forensic parameters of 57 autosomal InDels (A-InDels) included in our used new-generation InDel amplification system showed that this focused InDel panel is informative and polymorphic in Dingjie Sherpas, suggesting that it can be used as the supplementary tool for forensic personal identification and parentage testing in Dingjie Sherpas. Descriptive findings from the PCA, ADMIXTURE, and TreeMix-based phylogenies suggested that studied Nepal Sherpas showed excess allele sharing with neighboring Tibeto-Burman Tibetans. Furthermore, patterns of allele sharing in f-statistics demonstrated that Nepal Sherpas had a different evolutionary history compared with their neighbors from Nepal (Newar and Gurung) but showed genetic similarity with 2700-year-old Chokhopani and modern Tibet Tibetans. QpAdm/qpGraph-based admixture sources and models further showed that Sherpas, core Tibetans, and Chokhopani formed one clade, which could be fitted as having the main ancestry from late Neolithic Qijia millet farmers and other deep ancestries from early Asians. Chromosome painting profiles and shared IBD fragments inferred from fineSTRUCTURE and ChromoPainter not only confirmed the abovementioned genomic affinity patterns but also revealed the fine-scale genetic microstructures among Sino-Tibetan speakers. Finally, natural-selection signals revealed via iHS, nSL and iHH12 showed natural selection signatures associated with disease susceptibility in Sherpas. Generally, we provided the comprehensive landscape of admixture and evolutionary history of Sherpa people based on the shared alleles and haplotypes from the InDel-based genotype data and high-density genome-wide SNP data. The more detailed genetic landscape of Sherpa people should be further confirmed and characterized via ancient genomes or single-molecule real-time sequencing technology.

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17. The genetic structure and East-West population admixture in northwest China inferred from genome-wide array genotyping

Frontiers in Genetics, accepted 06 December 2021

Bin Ma, Jinwen Chen, Xiaomin Yang, Jingya Bai, Siwei Ouyang, Xiaodan Mo, Wangsheng Chen, Chuan-Chao Wang and  Xiangjun Hai 





Northwest China is a contacting region for East and West Eurasia and an important center for investigating the migration and admixture history of human populations. However, the comprehensive genetic structure and admixture history of the Altaic speaking populations and Hui group in Northwest China were still not fully characterized due to insufficient sampling and the lack of genome-wide data. Thus, We genotyped genome-wide SNPs for 140 individuals from 5 Chinese Mongolic, Turkic speaking groups including Dongxiang, Bonan, Yugur and Salar, as well as the Hui group. Analysis based on allele-sharing and haplotype-sharing were used to elucidate the population history of Northwest Chinese populations, including PCA, ADMIXTURE, pairwise Fst genetic distance, f-statistics, qpWave/qpAdm and ALDER, fineSTRUCTURE and GLOBETROTTER. We observed Dongxiang, Bonan, Yugur, Salar and Hui people were admixed populations deriving ancestry from both East and West Eurasians, with the proportions of West Eurasian related contributions ranging from 9% to 15%. The genetic admixture was probably driven by male-biased migration-showing a higher frequency of West Eurasian related Y chromosomal lineages than that of mtDNA detected in Northwest China. ALDER-based admixture and haplotype-based GLOBETROTTER showed this observed West Eurasian admixture signal was introduced into East Eurasia approximately 700~1000 years ago. Generally, our findings provided supporting evidence that the flourish transcontinental communication between East and West Eurasia played a vital role in the genetic formation of northwest Chinese populations.

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18. The locutor-referential pronoun in Zhoutun

Himalayan Linguistics, Vol. 20(3) 2021

Chenlei Zhou





This paper explores a special pronoun, the locutor-referential pronoun tha in Zhoutun, a Tibetanized Chinese variant spoken in the Amdo Sprachbund. Two rules of the use of tha are found in this paper. Rule 1: If tha occurs in a complement clause of a speech verb, it refers to the internal locutor. Rule 2: If tha occurs in an environment other than a complement clause of a speech verb, it refers to the narrative locutor. If only rule 1 is followed, then tha can be considered a logophoric pronoun; however, tha is special in that it can also be used in the context to which rule 2 applies, a usage that does not fit the definition of a logophoric pronoun. The use of tha is not obligatory. An inherited form from Mandarin Chinese, the formation of the locutor-referential tha has to do with the contact with Amdo Tibetan and its probable evolving pathway is “third-person pronoun> logophoric pronoun> locutor-referential pronoun”.

19. Intertwined model of syntactic borrowing in the Gansu-Qinghai linguistic area

Himalayan Linguistics, Vol. 20(3) 2021


Dan Xu





This paper studies two grammatical cases in the Gansu-Qinghai linguistic area. Accusative-dative, a syncretic case largely attested in Sinitic languages, is also found in Bao’an and Tu, even if in a very limited use. The Sinitic languages have acquired this syncretic case marking through pattern reduplication due to language contact, while Bao’an and Tu have this innovation owing to the internal mechanisms of their language. The second phenomenon concerns possessor constructions in which the subject-possessor must be marked by a dative case. This marking is seen in all non-Sinitic languages in the Gansu-Qinghai linguistic area and has begun to appear in Sinitic languages. Multiple paths for borrowing between and inside languages in this area present an intertwined model of language borrowing. Linxia City and its closest counties should be the spreading center of these new syntactic devices, and Muslim populations speaking different languages may form a spreading net.

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20. Genomic Insights Into the Population History and Biological Adaptation of Southwestern Chinese Hmong–Mien People

Frontiers in Genetics, 03 January 2022

doi: 10.3389/fgene.2021.815160

Yan Liu, Jie Xie, Mengge Wang, Changhui Liu, Jingrong Zhu, Xing Zou, Wenshan Li, Lin Wang, Cuo Leng, Quyi Xu, Hui-Yuan Yeh, Chuan-Chao Wang, Xiaohong Wen, Chao Liu and Guanglin He





Hmong–Mien (HM) -speaking populations, widely distributed in South China, the north of Thailand, Laos, and Vietnam, have experienced different settlement environments, dietary habits, and pathogenic exposure. However, their specific biological adaptation remained largely uncharacterized, which is important in the population evolutionary genetics and Trans-Omics for regional Precision Medicine. Besides, the origin and genetic diversity of HM people and their phylogenetic relationship with surrounding modern and ancient populations are also unknown. Here, we reported genome-wide SNPs in 52 representative Miao people and combined them with 144 HM people from 13 geographically representative populations to characterize the full genetic admixture and adaptive landscape of HM speakers. We found that obvious genetic substructures existed in geographically different HM populations; one localized in the HM clines, and others possessed affinity with Han Chinese. We also identified one new ancestral lineage specifically existed in HM people, which spatially distributed from Sichuan and Guizhou in the north to Thailand in the south. The sharing patterns of the newly identified homogenous ancestry component combined the estimated admixture times via the decay of linkage disequilibrium and haplotype sharing in GLOBETROTTER suggested that the modern HM-speaking populations originated from Southwest China and migrated southward in the historic period, which is consistent with the reconstructed phenomena of linguistic and archeological documents. Additionally, we identified specific adaptive signatures associated with several important human nervous system biological functions. Our pilot work emphasized the importance of anthropologically informed sampling and deeply genetic structure reconstruction via whole-genome sequencing in the next step in the deep Chinese Population Genomic Diversity Project (CPGDP), especially in the regions with rich ethnolinguistic diversity.

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21. Studying Han and non-Han Languages in Northern China from an Interdisciplinary Perspective



Chinese Social Sciences Today (中国社会科学报 ), 04 January 2022

Dan Xu

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22. Extensive ethnolinguistic diversity at the crossroads of North China and South Siberia reflects multiple sources of genetic diversity

Journal of Systematics and Evolution


09 January 2022

Guanglin He, Mengge Wang, Xing Zou, Hui-Yuan Yeh, Changhui Liu, Chao Liu, Gang Chen, Chuan-Chao Wang





North China and South Siberia, populated by Altaic- and Sino-Tibetan-speaking populations, possess extensive ethnolinguistic diversity and serve as the crossroads for the initial peopling of America and western-eastern trans-continental communication. However, the population genetic structure and admixture history of northern East Asians remain poorly understood due to a lack of genome-wide data, especially for Mongolic-speaking people in China. We genotyped genome-wide SNPs for 510 individuals from 38 Mongolic, Tungusic, and Sinitic speaking populations. We first explored the shared alleles and haplotypes within the studied groups. We then merged with 3508 published modern and ancient Eurasian individuals to reconstruct the deep evolutionary and natural selection history of northern East Asians. We identified genetic substructures within Altaic-speaking populations: Western Turkic people harboured more western Eurasian related ancestry; Northern Mongolic people in Siberia and eastern Tungusic people in Amur River Basin (ARB) possessed a majority of Neolithic ARB related ancestry; Southern Mongolic people in China possessed apparent genetic influence from Neolithic Yellow River Basin (YRB) farmers. Additionally, we found the differentiated admixture history between western and eastern Mongolians and geographically close Northeast Hans: the former received a genetic impact from western Eurasians, and the latter retained the primary Neolithic YRB and ARB ancestry. Moreover, we demonstrated that Kalmyk people from the northern Caucasus Mountain possessed a strong genetic affinity with Neolithic Mongolian Plateau (MP) people, supporting the hypothesis of their eastern Eurasian origin and long-distance migration history. We also illuminated that historical pastoral empires in the MP contributed considerably to the gene pool of northern Mongolic people but rarely to the southern ones. We finally found natural selection signatures in Mongolians associated with alcohol metabolism. Our results demonstrated that the Neolithic ancestral sources from the MP or ARB played an important role in spreading Altaic populations and languages. The observed multi-sources of genetic diversity contributed significantly to the extensive ethnolinguistic diversity in northern East Asia.

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23. Reconstructing the genetic admixture history of Tai-Kadai and Sinitic people:

Insights from genome-wide SNP data from South China

Journal of Systematics and Evolution


January 2022


doi: 10.1111/jse.12825

Mengge Wang, Guanglin He, Xing Zou, Pengyu Chen, Zheng Wang, Renkuan Tang, Xiaomin Yang, Jing Chen, Meiqing Yang, Yingxiang Li, Jing Liu, Fei Wang, Jing Zhao, Jianxin Guo, Rong Hu, Lan‐Hai Wei, Gang Chen, Hui‐Yuan Yeh, and  Chuan-Chao Wang





South China (SC) was a region with mixed rice–millet farming during the Middle Neolithic period and was also suggested to be the homeland of Tai‐Kadai (TK)‐speaking people. However, the formations of inland TK‐speaking people and southwestern Hans are far from clear due to very few studies on this subject. Here, we reveal the spatiotemporally demographic history of SC by analyzing newly‐generated genome‐wide SNP data of 115 modern southwestern individuals and find that inland TK‐speaking Dongs and Bouyeis have a close genomic affinity to coastal TK/Austronesian (AN)‐speaking people and Neolithic Yangtze River basin (YZRB) farmers, while southwestern Hans and TK‐speaking Gelaos possess a close genomic affinity to Neolithic Yellow River basin (YRB) farmers. Genetic differentiations are identified among TK people from SC and Southeast Asia, and between northern and southern inland Chinese TK people, in which the identified shared genetic ancestry between TK and AN people highlights a common origin of AN/TK groups. Conclusively, our findings indicate that millet farmers deriving from the YRB and rice farmers deriving from the YZRB substantially contribute to the present‐day inland TK speakers and southwestern Hans via a two‐way admixture scenario of bi‐directional gene‐flow events, which facilitates the formation of a modern two‐way genetic admixture profile.

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24. Fine-Scale Population Admixture Landscape of Tai–Kadai-Speaking Maonan in Southwest China Inferred From Genome-Wide SNP Data

Frontiers in Genetics


17 February 2022


Jing Chen, Guanglin He, Zheng Ren, Qiyan Wang, Yubo Liu, Hongling Zhang, Meiqing Yang, Han Zhang, Jingyan Ji, Jing Zhao, Jianxin Guo, Jinwen Chen, Kongyang Zhu, Xiaomin Yang, Rui Wang, Hao Ma, Le Tao, Yilan Liu, Qu Shen, Wenjiao Yang, Chuan-Chao Wang and Jiang Huang





Guizhou Province harbors extensive ethnolinguistic and cultural diversity with Sino- Tibetan-, Hmong–Mien-, and Tai–Kadai-speaking populations. However, previous genetic analyses mainly focused on the genetic admixture history of the former two linguistic groups. The admixture history of Tai–Kadai-speaking populations in Guizhou needed to be characterized further. Thus, we genotyped genome-wide SNP data from 41 Tai–Kadai-speaking Maonan people and made a comprehensive population genetic analysis to explore their genetic origin and admixture history based on the pattern of the sharing alleles and haplotypes. We found a genetic affinity among geographically different Tai–Kadai-speaking populations, especially for Guizhou Maonan people and reference Maonan from Guangxi. Furthermore, formal tests based on the f3/f4-statistics further identified an adjacent connection between Maonan and geographically adjacent Hmong–Mien and Sino-Tibetan people, which was consistent with their historically documented shared material culture (Zhang et al., iScience, 2020, 23, 101032). Fitted qpAdm-based two-way admixture models with ancestral sources from northern and southern East Asians demonstrated that Maonan people were an admixed population with primary ancestry related to Guangxi historical people and a minor proportion of ancestry from Northeast Asians, consistent with their linguistically supported southern China origin. Here, we presented the landscape of genetic structure and diversity of Maonan people and a simple demographic model for their evolutionary process. Further whole-genome- sequence–based projects can be presented with more detailed information about the population history and adaptative history of the Guizhou Maonan people.

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25. The genomic formation of Tanka people, an isolated “gypsies in water” in the coastal region of Southeast China

American Journal of Biological Anthropology


05 March 2022


Guanglin He, Yunhe Zhang, Lan-Hai Wei, Mengge Wang, Xiaomin Yang, Jianxin Guo, Rong Hu, Chuan-Chao Wang and Xianqing Zhang





Objectives: Three different hypotheses proposed via the controversial evidence from cultural, anthropological, and uniparental genetic analyses, respectively, stated that Tanka people probably originated from Han Chinese, ancient Baiyue tribe, or the admixture of them. Therefore, the genetic origin and admixture history of the Tanka people, an isolated “Gypsies in water” in the coastal region of Southeast China, are needed to be genetically clarified using genome-wide SNP data.

Materials and methods: To elucidate the genetic origin of the Southeast Tanka peo- ple and explore their genetic relationship with surrounding indigenous Tai-Kadai (TK), Hmong-Mien (HM), and Austronesian (AN) people and Neolithic-to-historic ancients from the Yellow River Basin (YRB) and Fujian, we conducted a large-scale population genomic study among 1498 modern and ancient Eurasians, in which 73 Tanka and 4 Han people were first reported here. Both allele-shared and haplotype-based statis- tical methods were used here, including PCA, ADMIXTURE, f-statistics, ALDER, qpGraph/TreeMix and qpAdm/qpWave, ChromoPainter, and fineSTRUCTURE.

Results: We found a specific genetic cline in PCA plots and detected the Tanka- specific homogeneous ancestry in model-based ADMIXTURE, suggesting differenti- ated demographic history between Tanka and surrounding Hans. Formal tests based on sharing allele patterns showed a close relationship between Tanka people and Han Chinese, but the Tanka population harbored more southern indigenous East Asian ancestry related to AA/HM/TK people compared with southern Hans. Besides, the reconstructed differentiated demographic history revealed that southern Xinshizhou Tankas harbored more ancestry related to the TK people or coastal ancient Neolithic to Bronze Age southern East Asians compared with northern Shacheng Tankas. The qpGraph/TreeMix-based phylogenetic framework, qpAdm/ qpWave-based admixture modeling among ancient northern and southern East Asians and fineSTRUCTURE-based dendrogram further demonstrated that the primary ancestry of modern Tankas derived from ancient millet farmers in the YRB with addi- tional admixture from multiple southern East Asian sources. 

Discussion: Sharing ancestry estimated from the f-statistics and sharing haplotypic landscape inferred from the ChromoPainter and fineSTRUCTURE showed that South- east Tanka people not only had a close genetic relationship with both northern Hans and YRB millet farmers but also possessed southern East Asian ancestry related to AA/HM/TK speakers. Our genomic data and fitted admixture models supported mod- ern Tanka originated from ancient North China and obtained additional gene flow from ancient southern East Asians in the processes of southward migrations. 

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26. Fine-Scale Genetic Profile and Admixture History of Two Hmong-Mien–Speaking Miao Tribes from Southwest China Inferred from Genome-wide Data

Human Biology, early views

Hui Tan, Rui Wang, and Chuan-Chao Wang





As the dominant indigenous minority in southern China, Hmong-Mien-speaking Miao people were thought to be the descendants of Neolithic Yangtze rice farmers. However, the fine-scale population structure and genetic profile of the Miao populations remain unclear due to the limited Miao samples from southern China and Southeast Asia. We genotyped 19 individuals from the two largest Miao tribes in Guizhou Province (Southwest China) via SNP chips and co-analyzed the data with published modern and ancient East Asians. The Guizhou Miao displayed a closer genomic affinity with present-day and Neolithic to Iron Age southern East Asians (SEAs) than with most northern East Asians (NEAs). The genetic substructure within Miao groups was driven by different levels of genetic interaction with other ethnolinguistic groups: Hunan Miao (Central China) harbored higher proportions of NEA-related ancestry; Guizhou Miao (Southwest China) and Vietnam Miao (mainland Southeast Asia) received additional gene flow mainly from surrounding groups with Tai-Kadai–related ancestry. There were also more complex admixture events in the newly studied groups between Guizhou Xijiang Miao and surrounding populations compared with Guizhou Congjiang Miao. The qpAdm model further demonstrated that the primary ancestry of Hunan Miao, Guizhou Miao studied here, and Vietnam Miao derived from ancient SEA-related ancestry (represented by coastal early Neolithic SEA Liangdao2), with the additional gene flow from ancient NEA-related ancestry (represented by spatiotemporally inland Yellow River farmers), with slightly different proportions. Our genomic evidence reveals the complex and distinct demographic history of different Miao tribes.

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27. Genetic Admixture History and Forensic Characteristics of Guizhou Sui People Inferred From Autosomal Insertion/Deletion and Genome-Wide Single-Nucleotide Polymorphisms

Frontiers in Ecology and Evolution 


10 may 2022



Meiqing Yang, Xiaomin Yang, Zheng Ren, Guanglin He, Han Zhang, Qiyang Wang, Yubo Liu, Hongling Zhang, Jingyan Ji, Jing Chen, Jianxin Guo, Jiang Huang and Chuan-Chao Wang





Insertion-deletion (Indel) serves as one of the important markers in forensic personal identification and parentage testing, especially for cases with degraded samples. However, the genetic diversity and forensic features in ethnolinguistically diverse southwestern Chinese populations remain to be explored. Sui, one Tai-Kadai- speaking population residing in Guizhou, has a complex genetic history based on linguistic, historic, and anthropological evidence. In this study, we genotyped 30 Indels from 511 Guizhou Sui individuals and obtained approximately 700,000 genome- wide single-nucleotide polymorphisms (SNPs) in 15 representative Sui individuals to comprehensively characterize the genetic diversity, forensic characteristics, and genomic landscape of Guizhou Sui people. The estimated forensic statistically allele frequency spectrum and parameters demonstrated that this Indels panel was polymorphic and informative in Tai-Kadai populations in southern China. Results of principal component analysis (PCA), STRUCTURE, and phylogenetic trees showed that Guizhou Sui had a close genetic relationship with geographically close Tai-Kadai and Hmong-Mien people. Furthermore, genomic analysis based on the Fst and f4-statistics further suggested the genetic affinity within southern Chinese Tai-Kadai-speaking populations and a close relationship with geographically adjoining Guizhou populations. Admixture models based on the ADMIXTURE, f4, three-way qpAdm, and ALDER results demonstrated the interaction between the common ancestor for Tai-Kadai/Austronesian, Hmong-Mien, and Austroasiatic speaking populations played a significant role in the formation of modern Tai-Kadai people. We observed a sex-biased influence in Sui people by finding that the dominant Y chromosomal type was a Hmong- Mien specific lineage O2a2a1a2a1a2-N5 but the mtDNA lineages were commonly found in Tai-Kadai populations. The additional southward expansion of millet farmers in the Yellow River Basin has impacted the gene pool of southern populations including Tai- Kadai. The whole-genome sequencing in the future will shed more light on the finer genetic profile of Guizhou populations.

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28. Genomic Insights Into the Unique Demographic History and Genetic Structure of Five Hmong-Mien-Speaking Miao and Yao Populations in Southwest China

Frontiers in Ecology and Evolution


09 June 2022


doi: 10.3389/fevo.2022.849195

Meiqing Yang, Guanglin He, Zheng Ren, Qiyan Wang, Yubo Liu,
Hongling Zhang, Han Zhang, Jing Chen, Jingyan Ji, Jing Zhao,
Jianxin Guo, Kongyang Zhu, Xiaomin Yang, Rui Wang, Hao Ma, Chuan-Chao Wang and Jiang Huang





Southern China was the original center of multiple ancestral populations related to modern Hmong-Mien, Tai-Kadai, Austroasiatic, and Austronesian people. More recent genetic surveys have focused on the fine-scale genetic structure and admixture history of southern Chinese populations, but the genetic formation and diversification of Hmong-Mien speakers are far from clear due to the sparse genetic sampling. Here, we reported nearly 700,000 single-nucleotide polymorphisms (SNPs) data from 130 Guizhou Miao and Yao individuals. We used principal component analysis, ADMIXTURE, f-statistics, qpAdm, phylogenetic tree, fineSTRUCTURE, and ALDER to explore the fine-scale population genetic structure and admixture pattern of Hmong-Mien people. The sharing allele patterns showed that our studied populations had a strong genetic affinity with ancient and modern groups from southern and southeastern East Asia. We identified one unique ancestry component maximized in Yao people, which widely existed in other Hmong-Mien-speaking populations in southern China and Southeast Asia and ancient samples of Guangxi. Guizhou Hmong-Mien speakers harbored the dominant proportions of ancestry related to southern indigenous East Asians and minor proportions of northern ancestry related to Yellow River farmers, suggesting the possibility of genetic admixture between Hmong-Mien people and recent southward Sino-Tibetan-related populations. Furthermore, we found a genetic substructure among geographically different Miao and Yao people in Leishan and Songtao. The Yao and Miao people in Leishan harbored more southern East Asian ancestry, but Miao in Songtao received more northern East Asian genetic influence. We observed high mtDNA but low Y-chromosome diversity in studied Hmong-Mien groups, supporting the role of sex- specific residence in influencing human genetic variation. Our data provide valuable clues for further exploring population dynamics in southern China.

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29. Genomic Insights Into the Demographic History of the Southern Chinese

Frontiers in Ecology and Evolution 


30 June 2022


doi: 10.3389/fevo.2022.853391

Xiufeng Huang, Zi-Yang Xia, Xiaoyun Bin, Guanglin He, Jianxin Guo, Atif Adnan, Lianfei Yin, Youyi Huang, Jing Zhao, Yidong Yang, Fuwei Ma, Yingxiang Li, Rong Hu, Tianle Yang, Lan-Hai Wei and Chuan-Chao Wang





Southern China is the birthplace of rice-cultivating agriculture and different language families and has also witnessed various human migrations that facilitated cultural diffusions. The fine-scale demographic history in situ that forms present-day local populations, however, remains unclear. To comprehensively cover the genetic diversity in East and Southeast Asia, we generated genome-wide SNP data from 211 present-day Southern Chinese and co-analyzed them with ∼1,200 ancient and modern genomes. In Southern China, language classification is significantly associated with genetic variation but with a different extent of predictability, and there is strong evidence for recent shared genetic history particularly in Hmong–Mien and Austronesian speakers. A geography- related genetic sub-structure that represents the major genetic variation in Southern East Asians is established pre-Holocene and its extremes are represented by Neolithic Fujianese and First Farmers in Mainland Southeast Asia. This sub-structure is largely reduced by admixture in ancient Southern Chinese since > ∼2,000 BP, which forms a “Southern Chinese Cluster” with a high level of genetic homogeneity. Further admixture characterizes the demographic history of the majority of Hmong–Mien speakers and some Kra-Dai speakers in Southwest China happened ∼1,500–1,000 BP, coeval to the reigns of local chiefdoms. In Yellow River Basin, we identify a connection of local populations to genetic sub-structure in Southern China with geographical correspondence appearing > ∼9,000 BP, while the gene flow likely closely related to “Southern Chinese Cluster” since the Longshan period (∼5,000–4,000 BP) forms ancestry profile of Han Chinese Cline.

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30. Multidisciplinary lines of evidence reveal East/Northeast Asian origins of agriculturalist/pastoralist residents at a Han dynasty military outpost in ancient Xinjiang

Frontiers in Ecology and Evolution


 10 August 2022



Edward Allen, Yao Yu, Xiaomin Yang, Yiran Xu, Panxin Du, Jianxue Xiong, Dian Chen, Xiaohong Tian, Yong Wu, Xiaoli Qin, Pengfei Sheng, Chuan-Chao Wang and Shaoqing Wen




Han/non-Han interactions were engrained among the border regions of ancient Imperial China. Yet, little is known about either the genetic origins or the lifeways of these border peoples. Our study applies tools from ancient deoxyribonucleic acid (DNA) and stable isotope analysis to the study of a Han dynasty population at the Shichengzi site in modern-day Xinjiang. Isotopic analysis (δ13C and δ15N) of human (n = 8), animal (n = 26), and crop remains (n= 23) from Shichengzi indicated that dietary patterns among site inhabitants could be split among agro-pastoral and agricultural groups based on differences in the collagen 15N ratios. DNA analysis divided the four Shichengzi samples into two groups, with one group primarily harboring the ancient Northeast Asian (ANA) related ancestry, while the other showed a dominant Late Neolithic Yellow River (YR_LN) related ancestry. Both ancient DNA and stable isotope evidence point to the Northeast Asian origins of pastoralists and East Asian origins of Han agriculturalists, who, nonetheless, shared a single burial space at Shichengzi. This study thus provides clear evidence for the multiple origins and identities of populations across the porous border represented by the Han Empire and surrounding regions and proposes a new model for the interpretation of border culture in early Imperial China.

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31. Genomic insight into the population history and biological adaptations of high-altitude Tibetan highlanders in Nagqu

Frontiers in Ecology and Evolution


17 August 2022



Lifeng Ma, Rui Wang, Siwei Feng, Xiaomin Yang, Jing Li, Zhiying Zhang, Haoshuang Zhan, Yao Wang, Ziyang Xia, Chuan-Chao Wang and Longli Kang




Tibetan, one of the largest indigenous populations living in the high-altitude region of the Tibetan Plateau (TP), has developed a suite of physiological adaptation strategies to cope with the extreme highland environment in TP. Here, we reported genome-wide SNP data from 48 Kham-speaking Nagqu Tibetans and analyzed it with published data from 1,067 individuals in 167 modern and ancient populations to characterize the detailed Tibetan subgroup history and population substructure. Overall, the patterns of allele sharing and haplotype sharing suggested (1) the relatively genetic homogeny between the studied Nagqu Tibetans and ancient Nepalese as well as present-day core Tibetans from Lhasa, Nagqu, and Shigatse; and (2) the close relationship between our studied Kham-speaking Nagqu Tibetans and Kham-speaking Chamdo Tibetans. The fitted qpAdm models showed that the studied Nagqu Tibetans could be fitted as having the main ancestry from late Neolithic upper Yellow River millet farmers and deeply diverged lineages from Southern East Asians (represented by Upper Paleolithic Guangxi_Longlin and Laos_Hoabinhian), and a non-neglectable western Steppe herder-related ancestry (∼3%). We further scanned the candidate genomic regions of natural selection for our newly generated Nagqu Tibetans and the published core Tibetans via FST, iHS, and XP-EHH tests. The genes overlapping with these regions were associated with essential human biological functions such as immune response, enzyme activity, signal transduction, skin development, and energy metabolism. Together, our results shed light on the admixture and evolutionary history of Nagqu Tibetan populations.

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32. The Shaping and Development of Huaxia (Chinese) Agriculture through the Book of Odes


中国农史 (Agricultural History of China)


04 2022

XU Dan



The Book of Odes represents the Chinese language from around the 11th to 6th centuries BC. This article attempts to compare the previous research results of the Book of Odes with recent archaeological discoveries to explore the maturation and development of China’s agriculture. The take-off of a civilization re­ quires material bases. The poems in the Book of Odes, especially the early ones, were collected during the flourishing period of the Chinese civilization, a period which also corresponds to the shaping and develop­ ment of Chinese agriculture. This epoch also matches the time of food globalization (Jones et al. 2011). This period is the stage where the Chinese fanning strides into the "prosperity of five crops" phase. The vocabulary- in the Book o f Odes reflects a typical farming one. I will compare the grain vocabulary in the Book with ar­ chaeological findings. Linguistics and archaeology studies can complement, corroborate or supplement each other. It can also correct some misunderstandings and misinterpretations in the study of the Book of Odes.

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33. Genetic substructure of Guizhou Tai-Kadai-speaking people inferred from genome-wide single nucleotide polymorphisms data

Frontiers in Ecology and Evolution


13 Septembre 2022



Zheng Ren, Meiqing Yang, Xiaoye Jin, Qiyan Wang, Yubo Liu, Hongling Zhang, Jingyan Ji, Chuan-Chao Wang and Jiang Huang




The genome-wide characteristics and admixture history of the Tai-Kadai- speaking populations are essential for understanding the population genetic diversity in southern China. We genotyped about 700,000 single nucleotide polymorphisms (SNPs) of 239 individuals from six Tai-Kadai-speaking populations residing in the mountainous Guizhou Province of southwestern China. We merged the genome-wide data with available populations and ancients in East and Southeast Asia to infer Tai-Kadai-speaking populations’ admixture history and genetic structure. We observed a genetic substructure within the studied six populations in the PCA, ADMIXTURE, ChromoPainter, GLOBETROTTER, f-statistics, and qpWave analysis. The Dong, Zhuang, and Bouyei people had a strong genetic affinity with other Tai-Kadai-speaking and Austronesian groups in the surrounding area. However, Gelao showed an affinity to Sino-Tibetan groups, and Mulao people were genetically close to Hmong-Mien populations. qpAdm further illuminated that Gelao and Dong_Tongren composited more Han-related ancestry than Dong, Zhuang, Bouyei, and Mulao people. Meanwhile, we observed high frequencies of Y-chromosome haplogroup O in studied Tai-Kadai-speaking groups except for Gelao people with a high haplogroup N frequency. From the maternal side, haplogroup M7 was frequent in studied populations except for Tongren Dong, who had a high frequency of haplogroup B5. Our newly reported data are helpful for further exploring population dynamics in southern China.

33. Genetic substructure of Guizhou Tai-Kadai-speaking people inferred from genome-wide single nucleotide polymorphisms data

Frontiers in Ecology and Evolution


13 Septembre 2022



Zheng Ren, Meiqing Yang, Xiaoye Jin, Qiyan Wang, Yubo Liu, Hongling Zhang, Jingyan Ji, Chuan-Chao Wang and Jiang Huang




The genome-wide characteristics and admixture history of the Tai-Kadai- speaking populations are essential for understanding the population genetic diversity in southern China. We genotyped about 700,000 single nucleotide polymorphisms (SNPs) of 239 individuals from six Tai-Kadai-speaking populations residing in the mountainous Guizhou Province of southwestern China. We merged the genome-wide data with available populations and ancients in East and Southeast Asia to infer Tai-Kadai-speaking populations’ admixture history and genetic structure. We observed a genetic substructure within the studied six populations in the PCA, ADMIXTURE, ChromoPainter, GLOBETROTTER, f-statistics, and qpWave analysis. The Dong, Zhuang, and Bouyei people had a strong genetic affinity with other Tai-Kadai-speaking and Austronesian groups in the surrounding area. However, Gelao showed an affinity to Sino-Tibetan groups, and Mulao people were genetically close to Hmong-Mien populations. qpAdm further illuminated that Gelao and Dong_Tongren composited more Han-related ancestry than Dong, Zhuang, Bouyei, and Mulao people. Meanwhile, we observed high frequencies of Y-chromosome haplogroup O in studied Tai-Kadai-speaking groups except for Gelao people with a high haplogroup N frequency. From the maternal side, haplogroup M7 was frequent in studied populations except for Tongren Dong, who had a high frequency of haplogroup B5. Our newly reported data are helpful for further exploring population dynamics in southern China.

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34. Cultural and demic co-diffusion of Tubo Empire on Tibetan Plateau



Accepted 17 November 2022

Kongyang Zhu, Panxin Du, Jiyuan Li, Jianlin Zhang, Xiaojun Hu, Hailiang Meng, Liang Chen, Boyan Zhou, Xiaomin Yang, Jianxue Xiong, Edward Allen, Xiaoying Ren, Yi Ding, Yiran Xu, Xin Chang, Guanghui Dong, Chuan-Chao Wang, Shaoqing Wen




A high point of Tibetan Plateau (TP) civilization, the expansive Tubo Empire (618–842 AD) wielded great influence across ancient western China. However, whether the Tubo expansion was cultural or demic remains unclear due to sparse ancient DNA sampling. Here we reported ten ancient genomes at 0.017- to 0.867-fold coverages from the Dulan site with typical Tubo archaeological culture dating to 1308-1130 BP. Nine individuals from three different grave types have close relationship with previously reported ancient highlanders from the southwestern Himalayas and modern core-Tibetan populations. A Dulan-related Tubo ancestry contributed overwhelmingly (95%-100%) to the formation of modern Tibetans. A genetic outlier with dominant Eurasian steppe-related ancestry suggesting a potential population movement into the Tubo-controlled regions from Central Asia. Together with archeological evidence from burial styles and customs, our study suggested the impact of the Tubo empire on the northeast edge of the TP involved both cultural and demic diffusion.

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