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Whole-sequence analysis indicates that the Y chromosome C2*-Star Cluster traces back to ordinary Mongols, rather than Genghis Khan
Open accessAbstract
The Y-chromosome haplogroup C3*-Star Cluster (revised to C2*-ST in this study) was proposed to be the Y-profile of Genghis Khan. Here, we re-examined the origin of C2*-ST and its associations with Genghis Khan and Mongol populations. We analyzed 34 Y-chromosome sequences of haplogroup C2*-ST and its most closely related lineage. We redefined this paternal lineage as C2b1a3a1-F3796 and generated a highly revised phylogenetic tree of the haplogroup, including 36 sub-lineages and 265 non-private Y-chromosome variants. We performed a comprehensive analysis and age estimation of this lineage in eastern Eurasia, including 18,210 individuals from 292 populations. We discovered that the origin of populations with high frequencies of C2*-ST can be traced to either an ancient Niru'un Mongol clan or ordinary Mongol tribes. Importantly, the age of the most recent common ancestor of C2*-ST (2576 years, 95% CI = 1975-3178) and its sub-lineages, and their expansion patterns, are consistent with the diffusion of all Mongolic-speaking populations, rather than Genghis Khan himself or his close male relatives. We concluded that haplogroup C2*-ST is one of the founder paternal lineages of all Mongolic-speaking populations, and direct evidence of an association between C2*-ST and Genghis Khan has yet to be discovered.
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TL;DR
Whole-sequence and Y-STR analyses of the Y-chromosome lineage C2*-ST (redefined as C2b1a3a1-F3796) across eastern Eurasia indicate it originated ~2,576 years ago and expanded with Mongolic-speaking populations, not with Genghis Khan. The authors conclude C2*-ST is a founder paternal lineage of ordinary Mongol tribes, and direct evidence linking it to Genghis Khan has yet to be discovered.
Key findings
- Among 18,210 individuals from 292 populations, the authors identified 809 C2*-ST samples with 685 Y-STR haplotypes; the highest frequencies were in several Kazakh populations, followed by Mongolian, Buryat-Bargut, and Uzbek populations, while C2*-ST was absent or very low in non-Altaic populations except Hazara.
- The revised phylogeny of C2b1a3a1-F3796 contained 36 sublineages and 265 non-private Y-chromosome variants; its closest branch was C2b1a3a2-F8951 (the C2*-Daur clade).
- The TMRCA of C2*-ST estimated by BEAST with bModelTest was 2,576 years (95% CI = 1,975–3,178), which is older than the earliest Chinese historical record of Mongol tribes (~1,300 years ago) and consistent with the diffusion of all Mongolic-speaking populations rather than Genghis Khan.
- No donors carrying C2*-ST could trace their genealogy to Genghis Khan, and no self-claimed direct descendants of Genghis Khan carried C2*-ST; a documented direct descendant, Dayan Khan, had haplogroup C2c1a1a1-M407.
Methods
- Blood or saliva samples from 6,348 unrelated healthy males from 74 populations in eastern Eurasia were collected between 2005 and 2014; these were combined with literature data for a total of 18,210 individuals from 292 populations. Samples were genotyped for Y-chromosome markers and 17 Y-STR loci, and 17 selected C2*-ST samples were sequenced using Illumina HiSeq2000 with a ~11 M bp Y-chromosome capture, plus 17 previously published Y-chromosome sequences.
- The authors revised the phylogenetic tree following Y Chromosome Consortium guidelines, constructed median-joining Y-STR networks, plotted population frequencies geographically, and estimated coalescence times using ASD, BATWING, and BEAST v.2.4.3 with bModelTest and GTR+G+I models. Calibration points were haplogroup CT-M168 (71,760 years, 95% CI = 69,777–73,799) and the Aisin Gioro family split (470 ± 20 years ago), with a generation time of 30 years.
Limitations
- The paper states that direct genotyping of more well-documented male descendants of Genghis Khan from a wider geographic region is needed to definitively characterize the Y chromosome of Genghis Khan.
- The paper states that ASD may underestimate ages owing to the large number of identical Y-STR haplotypes in C2*-ST, and that BATWING produced older estimates than the other methods; the authors therefore preferred the BEAST whole-sequence estimates.
- Evident from the text: the study relies on modern population samples and historical records rather than ancient DNA or direct testing of Genghis Khan himself, and some subclade age estimates are based on small numbers of Y-chromosome sequences (e.g., C2b1a3a2-F8951, N = 3; C2b1a3a1c2a-F8949, N = 3).
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19 Own result · 15 Own interpretation · 12 Not checked yet · 4 Check pending · 4 Own method · 3 Sources not found · 3 No full text of the sources · 1 No basis given
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In the paper
“We observed the highest frequencies of C2*-ST in several Kazakh populations in Southeast Kazakhstan and Northwest China, followed by Mongolian, Buryat-Bargut, and Uzbeks.”
§ Whole-sequence analysis indicates that the Y chromosome C2*-Star Cluster traces back to ordinary Mongols, rather than Genghis > Results
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In the paper
“Y-chromosome sequence data also indicated that samples with DYS448 = 23 form a special sub-lineage, C2b1a3a1c2a-F8949 (Fig. 3 and Supplementary Table S3).”
§ Whole-sequence analysis indicates that the Y chromosome C2*-Star Cluster traces back to ordinary Mongols, rather than Genghis > Results
hedgedstates an association- Asserted: passes — The authors put it forward as true.
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In the paper
“Among 18,210 samples from 292 populations throughout eastern Eurasia, we identified 809 samples with 685 Y-STR haplotypes as C2*-ST (see Supplementary Table S2).”
§ Whole-sequence analysis indicates that the Y chromosome C2*-Star Cluster traces back to ordinary Mongols, rather than Genghis > Results
plainn = 809- Asserted: passes — The authors put it forward as true.
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In the paper
“By contrast, C2*-ST was absent or found at very low frequencies in non-Altaic populations.The only exceptions were Hazara.”
§ Whole-sequence analysis indicates that the Y chromosome C2*-Star Cluster traces back to ordinary Mongols, rather than Genghis > Results
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“As shown in the Y-STR network, a special branch with DYS448 = 23 (defined as the C2*-ST-Kazakh clade, Fig. 2 and Supplementary Table S2) emerged in the Kazakh-Kerey-Abakh population [7].”
§ Whole-sequence analysis indicates that the Y chromosome C2*-Star Cluster traces back to ordinary Mongols, rather than Genghis > Results
What the sources say
- Not checked yetcites [7] Abilev S, Malyarchuk B, Derenko M, Wozniak M, Grzybowski T, Zakharov I. The Y-chromosome C3* star-cluster attributed to Genghis Khan’s descendants is present at high frequency in the Kerey clan from Kazakhstan. Hum Biol. 2012;84:79–89. source
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Claims other papers make about this one
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In the citing paper
“Like all of the samples in the C3* tree (Fig. 2B), Kenggir males had the derived allele T for the marker F3796, which defines C3* Y-chromosomes (32), along with the upstream F1918 and other markers.”
From Genomes of the Golden Horde elites and their implications for the rulers of the Mongol Empire (2026) · cites this paper as [32]
In the citing paper
“Populations or individuals who claim descent from Genghis Khan had various paternal lineages (16, 32, 46, 47), and no consensus has been reached.”
From Genomes of the Golden Horde elites and their implications for the rulers of the Mongol Empire (2026) · cites this paper as [32]
In the citing paper
“The age estimation of modern east Eurasia Y-chromosome STR data belonging to the C2a1a3-F1918 was determined to be 6327 ± 1736 years (Wei et al., 2018).”
From Y chromosome evidence confirms northeast Asian origin of Xinjiang Kazakhs and genetic influence from 18th century expansion of Kerey clan (2023) · cites this paper as Wei et al., 2018
In the citing paper
“Based on these findings, we propose that haplogroup C2a1a3-F1918 originated in ancient northeast Asia and expanded to the Mongolia Plateau, later spreading westward to Central Asia and Europe, following the route of its subbranch C2a1a3a-F3796 (Wei et al., 2018).”
From Y chromosome evidence confirms northeast Asian origin of Xinjiang Kazakhs and genetic influence from 18th century expansion of Kerey clan (2023) · cites this paper as Wei et al., 2018
In the citing paper
“The dominant haplogroup C2a1a3-F1918, which originated in northeast Asia during the Neolithic Age, accounts for nearly half of the Altay and Ili Kazakhs.”
From Y chromosome evidence confirms northeast Asian origin of Xinjiang Kazakhs and genetic influence from 18th century expansion of Kerey clan (2023) · cites this paper as Wei et al., 2018
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In the paper
“Another 17 previously published Y-chromosome sequences were also used to construct a phylogeny of haplogroup C2*-ST, including one sequence from Yan et al. [12], three from Wei et al. [17], one from Karmin et al. [18], and 11 from Lippold et al. [19]”
§ Whole-sequence analysis indicates that the Y chromosome C2*-Star Cluster traces back to ordinary Mongols, rather than Genghis > Molecular methods
Reported 17 · recomputed 16
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Metadata
- Authors
- Wei, L.-H., Yan, S., Lu, Y., Wen, S.-Q., Huang, Y.-Z., Wang, L.-X., Li, S.-L., Yang, Y.-J., Wang, X.-F., Zhang, C., Xu, S.-H., Yao, D.-L., Jin, L., & Li, H.
- Affiliations
- Institut National des Langues et Civilisations Orientales (FR)
- Fudan University (CN)
- Max Planck Society (DE)
- Corresponding author
- Li, Hui
- Venue
- European Journal of Human Genetics
- Published
- 2018-01-22
- Volume
- 26
- Issue
- 2
- Pages
- 230-237
- ISSN
- 1018-4813 (ISSN-L)
- 1476-5438
- PMC
- PMC5839053
- OpenAlex
- W2790037793
- Semantic Scholar
- 5849a9e5bb3120d1e9e0e1be3a751cafb6c967cc
- Cited by
- 57
- References
- 34
- Citations by year
- 2026: 3 · 2025: 4 · 2024: 9 · 2023: 6 · 2022: 4 · 2021: 9 · 2020: 15 · 2019: 4 · 2018: 3
- FWCI
- 4.06
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- www.springer.com/tdm (tdm)
- Locations
- European Journal of Human Genetics (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (publishedVersion)
- Funders
- National Natural Science Foundation of China (31222030, 31671297, 31501011, 2016YFC0900300, 91331204, 31401060, 31271338)
- Science and Technology Commission of Shanghai Municipality (14YF1406800)
- Ministry of Education, India
- China Scholarship Council
- 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
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic Associations and Epidemiology
- Keywords
- Genghis Khan, Mongol expansion
- MeSH terms
- Haplotypes, Humans, Male, Founder Effect, Evolution, Molecular, Chromosomes, Human, Y, Asian People
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