Ancient genome of the Chinese Emperor Wu of Northern Zhou

[Shuzam87]

https://www.sciencedirect.com/science/ar...via%3Dihub

Ancient genome of the Chinese Emperor Wu of Northern Zhou

https://doi.org/10.1016/j.cub.2024.02.059

Highlights

• Wudi had a typical East or Northeast Asian facial appearance
  
• Pathogenic SNPs suggest an increased susceptibility of Wudi to stroke
  
• Wudi derived ancestry from Northeast Asians but also has Han-related admixture
  

Summary
Emperor Wu (武帝, Wudi) of the Xianbei-led Northern Zhou dynasty, named Yuwen Yong (宇文邕, 543–578 CE), was a highly influential emperor who reformed the system of regional troops, pacified the Turks, and unified the northern part of the country. His genetic profile and physical characteristics, including his appearance and potential diseases, have garnered significant interest from the academic community and the public. In this study, we have successfully generated a 0.343×-coverage genome of Wudi with 1,011,419 single-nucleotide polymorphisms (SNPs) on the 1240k panel. By analyzing pigmentation-relevant SNPs and conducting cranial CT-based facial reconstruction, we have determined that Wudi possessed a typical East or Northeast Asian appearance. Furthermore, pathogenic SNPs suggest Wudi faced an increased susceptibility to certain diseases, such as stroke. Wudi shared the closest genetic relationship with ancient Khitan and Heishui Mohe samples and modern Daur and Mongolian populations but also showed additional affinity with Yellow River (YR) farmers. We estimated that Wudi derived 61% of his ancestry from ancient Northeast Asians (ANAs) and nearly one-third from YR farmer-related groups. This can likely be attributed to continuous intermarriage between Xianbei royal families, and local Han aristocrats.1,2 Furthermore, our study has revealed genetic diversities among available ancient Xianbei individuals from different regions, suggesting that the formation of the Xianbei was a dynamic process influenced by admixture with surrounding populations.

This individual was determined as male, and his Y chromosome was assigned as a haplogroup C2a1a1b1a-F3830+, F8497− (Data S1A) by PCR-based targeted amplification covering 485 Y chromosome SNPs, which was further confirmed by hybridization capture data as a downstream haplogroup C2a1a1b1a2a1-FGC28857 × (FGC28846, Z44095, FGC31362, Z45818) (Data S1B). Due to the higher copy number of mtDNA versus nuclear DNA,14 we generated an mtDNA genome with coverage of 507.08 X and determined his haplogroup as C4a1a + 195.15,16 His paternal and maternal lineages can both be traced back to Northeast Asia and still reach moderate frequency at present.17,18,19

[Shuzam87]

Updated Version of Emperor Wu of Northern Zhou's data:

https://ngdc.cncb.ac.cn/gsa-human/browse/HRA006585

[ilabv]

Updated Version of Emperor Wu of Northern Zhou's data:

https://ngdc.cncb.ac.cn/gsa-human/browse/HRA006585

Send to Davidski

[Shuzam87]

Updated Version of Emperor Wu of Northern Zhou's data:

https://ngdc.cncb.ac.cn/gsa-human/browse/HRA006585

Newly updated YDNA: C-FGC28857 Mtdna: C4a1a-a5

https://www.theytree.com/sample/ea51f944...82bb6.html

[qijia]

I think western Zhou genome is more important. Their royals had N-M128, so what language did they speak for 730 years?

"According to the "Research Report on Ancient DNA of the Remains of the Western Zhou Cemetery in Xinancheng of the Longzi" released by it, 4 of the 7 male samples were tested for N-M128 (upstream of N-F1998), and the other three cases were: 1 Q1a-Y642, 1 C2a1-CTS2428, 1 N1b2e-M1846."

[Vinitharya]

I find it intriguing that of the male haplogroups in that study, none are O, which is by far the majority haplogroup of the Chinese.

[okarinaofsteiner]

I find it intriguing that of the male haplogroups in that study, none are O, which is by far the majority haplogroup of the Chinese.

This isn't surprising. The ruling family studied here was of "northern barbarian" lineage, and genetically distinct from both ancient and modern-day Northern Han Chinese.

I find it more interesting that several prominent figures in Ancient China (pre-imperial and early imperial) also didn't have Y-DNA haplogroup O. Confucius is alleged to have been haplogroup Q1a1a or C2.

Descendant testing of very ancient people has often proved quite unreliable. For example, over 1000 presumed paternal descendants of Confucius had their Y-DNA tested, and were found to belong to haplogroup C2 (c-M217) (46.06%), haplogroups Q1a1a1 (Q-M120) (27.01%), haplogroups O2 (O-M122) (20.66%), and other haplogroups (6.27%). Notwithstanding, both C2 and Q1a1a are relatively rare in China (about 10% and 0.5% respectively), so such high percentages indicate that one of them could really be the right lineage, with a higher statistical probability for Q1a1a. The TMRCA of those descendants should be compared. If one lineage dates back more or less to 2500 years ago, then it might be it.

[Ebizur]

I find it intriguing that of the male haplogroups in that study, none are O, which is by far the majority haplogroup of the Chinese.

This isn't surprising. The ruling family studied here was of "northern barbarian" lineage, and genetically distinct from both ancient and modern-day Northern Han Chinese.

Emperor Wu of Northern Zhou (北周武帝), who is the topic of the OP in this thread, is of known "northern barbarian" (in his case, Xianbei) lineage. He lived during the sixth century CE.

On the other hand, the specimens from the Xinancheng Cemetery (located in Zhangzi county, Changzhi prefecture-level city, Shanxi province) to which qijia and Vinitharya have referred in this thread represent members of the upper class of the Western Zhou (i.e. following the Shang and preceding the Eastern Zhou), and they have lived during the early first millennium BCE, more than a thousand years before Emperor Wu of Northern Zhou.

It surely must be quite surprising that the Ji (姬) clan of (Western) Zhou may have belonged to haplogroup N-M128, which would mean that they may be patrilineally more closely related to present-day members of N-P43 (including many people who currently speak a Uralic language or a Turkic language) than to most present-day Chinese people.

I find it more interesting that several prominent figures in Ancient China (pre-imperial and early imperial) also didn't have Y-DNA haplogroup O. Confucius is alleged to have been haplogroup Q1a1a or C2.

Descendant testing of very ancient people has often proved quite unreliable. For example, over 1000 presumed paternal descendants of Confucius had their Y-DNA tested, and were found to belong to haplogroup C2 (c-M217) (46.06%), haplogroups Q1a1a1 (Q-M120) (27.01%), haplogroups O2 (O-M122) (20.66%), and other haplogroups (6.27%). Notwithstanding, both C2 and Q1a1a are relatively rare in China (about 10% and 0.5% respectively), so such high percentages indicate that one of them could really be the right lineage, with a higher statistical probability for Q1a1a. The TMRCA of those descendants should be compared. If one lineage dates back more or less to 2500 years ago, then it might be it.

Q-M120 accounts for much more than 0.5% of present-day Chinese males. Perhaps that may be a typographical error for 2.5%.

According to official Chinese historical texts, Confucius should be a patrilineal descendant of the ruling family of the Shang, who preceded the aforementioned Western Zhou, via the Dukes of Song. Song, most of whose territory is now located in the eastern parts of Henan province, was one of many states under Zhou suzerainty; in essence, Song was a successor state of Shang, maintaining Shang traditions and lineage after the ascendance of Zhou. Song was located east of the original Zhou heartland as well as southeast of the Xinancheng Cemetery of Western Zhou; however, by the lifetime of Confucius, the center of Zhou power had shifted eastward (hence the historiographic distinction between the Western Zhou and the subsequent Eastern Zhou; Confucius lived during the time of the Eastern Zhou, about five centuries after his alleged Shang ancestors had capitulated to Zhou and about two centuries after the eastward relocation of the Zhou capital).

[CLTVTE]

As for yDNA Q-L472, which is an ancestor of both yDNA Q-L804 and yDNA Q-M120, "Bronze and Iron Age population movements underlie Xinjiang population history" reported an Iron Age sample from Xinjiang, which was determined to belong to either yDNA CT or yDNA Q1a (that is, Q-L472 in the version of yDNA nomenclature, specified in the article). This ancient specimen was reported to have a genetic affinity to the Bactria Margiana Archaeological complex’s specimens in the same article. This ancient specimen had mtDNA H14b2, which shares a mutation C7610T with the mtDNA M7c1c3am of the Austronesians of the Madagascar Island (https://www.yfull.com/mtree/M7c1c3am/). Consequently, the question may appear, which sort of yDNA CT affinity this ancient yDNA Q-L472-related specimen had. In “Human genetic history on the Tibetan Plateau in the past 5100 years”, yDNA D-M174 was also only determined as yDNA CT

C1258 L3085 Liushui Yutian County, Hetian Region 36,46 81,95 2433 South LSH_M50_5 LSH_IA3 LSH_IA3_oBMAC Included Iron Age - Teeth SNP_Capture DS_half 2992-2798 556 0,004 0 330 4,4 0,04 45957 mtDNA H14b2 M Q1 CT Q1 yDNA Q1a

In the article, co-authored by an American-born researcher and a Portuguese-Indian researcher (“Novel 4,400-year-old ancestral component in a tribe speaking a Dravidian language”), the Japanese (“JPN”) were used as an outgroup for the model, describing the ancestry, which was distributing between various Dravidian individuals (Supplementary Fig. 3), including the Koraga individuals, while the article stated that "the Elamo-Dravidian theory and the linguistic phylogeny of the Dravidian family tree provide chronological fits for the genetic findings presented here". In “Human genetic history on the Tibetan Plateau in the past 5100 years”, the ancient Qihe3 specimen of the Fujian Province acquired the BMAC-related Gonur1_BA component, while the article “Novel 4,400-year-old ancestral component in a tribe speaking a Dravidian language” reiterated the existence of the Western Eurasian-related “Ancestral North Indian (ANI)–Ancestral South Indian (ASI)–Ancestral Austroasiatic (AAA)” cline. However, in “The first maternal genetic study of hunter-gatherers from Vietnam”, authored by researchers from the Vietnam Academy of Science and Technology, the mtDNA of the ancient Qihe specimen clustered as an outgroup to mtDNAs of all used Austroasiatic individuals. Similarly, hypotheses on the relationship between the Elamite language and Austroasiatic languages are quite marginal and are not generally accepted hypotheses so far.

C3331 L6145 G218 Nileke County, Yili Region 43,8 82,51 1200 West G218_M1_Kin G218_IA_K G218_IA_K Not included Iron Age Scythian, Yuezhi, or Wusun-related[96-97] Teeth SNP_Capture DS_half 2336-1942* 21 0,002 0,1 177 8,9 0,04 41051 L6145_2d._L6236,L6145_2d._L6250 HV18 NLKG218_M1 M Q1b F no call
Additionally, an ancient sample C3331 from Xinjiang, having an yDNA, equivalent to yDNA Q-L56, which is closer related to Native Americans, was determined as yDNA F in "Bronze and Iron Age population movements underlie Xinjiang population history", while yDNA F as such was also reported from Tibet and Southern East Asia. This ancient sample did not show an affinity to the Bactria Margiana Archaeological complex’s specimens, but instead of this, this ancient sample had mtDNA R0>HV18+T16189C!, which is observed in Mesopotamian Arabs https://www.yfull.com/mtree/HV18/ The scope of settlement of yDNA Q-L56 individuals in the Near East remains to be clarified.

[sg_jun]

I find it intriguing that of the male haplogroups in that study, none are O, which is by far the majority haplogroup of the Chinese.

Y-Hg O likely only became the majority among Chinese during the late Warring States, Spring & Autumn period as well as Qin-Han dynasties; some of the largest lineages include O-F254, O-F656 truly underwent a population boom during this time and it is believed these 2 may represent the royal lineages of Han Dynasty and Yue kingdom respectively. Chu kingdom which was the second most powerful and largest state during the Warring States period was also centered around southern China where Hg O had a much larger population to begin with. In addition many southern Chinese do actually have Yue lineages (indigenous people of southern China related to SE Asians). Among most Southeast Asian ethnicities today haplogroup O is also by far the most populous (something like 80-90%).

[Ebizur]

I find it intriguing that of the male haplogroups in that study, none are O, which is by far the majority haplogroup of the Chinese.

Y-Hg O likely only became the majority among Chinese during the late Warring States, Spring & Autumn period as well as Qin-Han dynasties; some of the largest lineages include O-F254, O-F656 truly underwent a population boom during this time and it is believed these 2 may represent the royal lineages of Han Dynasty and Yue kingdom respectively. Chu kingdom which was the second most powerful and largest state during the Warring States period was also centered around southern China where Hg O had a much larger population to begin with. In addition many southern Chinese do actually have Yue lineages (indigenous people of southern China related to SE Asians). Among most Southeast Asian ethnicities today haplogroup O is also by far the most populous (something like 80-90%).

As I recall, every male specimen from the Taosi site for which Y-DNA results have been published to date has belonged to haplogroup O2-M122. Taosi is a site located in Xiangfen County, Linfen prefecture-level city, Shanxi Province and bearing remains attributed to the Longshan culture; the Taosi site is geographically quite close to the Xinancheng site of the Western Zhou mentioned previously in this thread.

Specimens from the Wanggou site in Henan Province, attributed to the Yangshao culture and associated with early silk fabrics, likewise have been assigned to Y-DNA haplogroup O, including a case of haplogroup O1b1a2c in addition to a case of haplogroup O2a2b1a1a1e2.

Members of O2a2b1a1 also have been found among specimens attributed to the Hongshan culture in Liaoning (Niuheliang site, Banlashan site, etc.).

So, at least haplogroup O2 has been present among populations in Shanxi, Henan, and Liaoning since the Neolithic.

[CLTVTE]

It has long been claimed by the known Japanese DNA website that the yDNA N-M128 is an yDNA of the local family of Shinto priests in Nanao City (七尾市) of the Ishikawa Prefecture of Japan, who have been taking care of the shine, having the word “Arakashihiko” in the shrine’s name. It appears that the surname of the family, taking care of the Arakashihiko-related shrine is “Shimizu” (清水), and this family is engaged in taking care of 26 other Shinto shrines in the same locality. Other men from Nanao, having the surname “Shimizu”, include a member of Nanao’s local government, who belongs to the ruling Liberal Democratic Party of Japan, as well as a young Japanese military man from Nanao, whose death during the attack on Pearl Harbor was mentioned by the “Mainichi Shimbun” newspaper. It should be mentioned that, according to the environmental data, the word “shimizu” (清水), meaning “clear water”, can appear as a place name wherever there is a beneficial source of underground water, emitting from the ground, even including a hot water spring. It means that, since “Shimizu” (清水) is the 18th-20th most common Japanese surname, which can have different ways of origin, totally unrelated Shimizu individuals should not apply the above and below information to themselves.

Judging by the yfull data, where there used to be slightly more than 100 Japanese male individuals, of whom 4 Japanese male individuals belonged to yDNA N-M128, there was an affinity to report yDNA N-M128 from the Japanese male population on the yfull site. In regard to the role of yDNA N-M128-related population, which would be relevant for the Japanese individuals, the article “The deep population history of northern East Asia from the Late Pleistocene to the Holocene” showed the following information: on the PCA, the yDNA N-M128 individual, being geographically assigned to North China, nonetheless, autosomally clustered very close to more southern specimens from the more rice-farming Jiangsu and Anhui Provinces of China (while it is known from the Japanese sources that at least some Japanese individuals, having the surname “Shimizu” (清水) and living in the region of Nanao City, Ishikawa Prefecture, had a knowledge of rice farming in the past), and the same yDNA N-M128 individual participated in a genetic cline, involving an yDNA O1b2-L682/mtDNA F1a1b Japanese specimen and an mtDNA D4a4a/yDNA O-F16241 (https://www.yfull.com/tree/O-F16241/ ) Japanese specimen, who has multiple Han Chinese relatives in China (see yDNA O-M134>O-F450>O-F122>O-F114>O-F629>O-F79>O-F46>O-F502>O-FGC85750>O-FGC16847>O-Z26092>O-F242>O-F1116>O-F273>O-Y20147>O-Z26101>O-Z26106>O-CTS4266>O-F14839>O-F23272 https://www.theytree.com/tree/O-MF174853), but the TMRCA of this yDNA O- F16241 subbranch is 3900 years on the yfull site and 3940 years on theytree.com site (https://www.theytree.com/tree/O-F17584), which is quite similar to the TMRCA of yDNA N-M128 (yDNA N-F710 https://www.yfull.com/tree/N-F710/).

According to some Japanese researchers, the element ara- in the “Arakashihiko” part of the name of the Arakashihiko-related shrine of Nanao City does not point to the origin from the ancient Korean kingdom of Silla, but points to the origin from the geographic territory of the iron-making Ara city-state kingdom of the Gaya(Kara) Confederacy (Gaya’s Japonic or Koreanic affinity is disputed), and Ara and the interaction with Ara’s ruler was mentioned in “Nihon Shoki”. The Arakashihiko-related shrine is located not far from the river named “Kumaki”, however, the full name of the shrine, cited by all available Japanese researchers, has a much more poetically sounding first component, though distantly resembling the name of the Kumaki river: Kumakabuto Arakashihiko (久麻加夫都阿良加志比古). The “Kuma Kabuto” (久麻加夫都) element may refer to a samurai helmet (kabuto), which is enforced or tightly decorated with the hair of a bear. The hieroglyphs 久麻 denote “a bear” (*kuma), and the form *kuma was claimed by Western researchers to be similar to *kuma, a word for “a bear” in one of the languages of the Kingdom of Paekche, whose affiliation to a Koreanic language or a Japonic language is disputed. Moreover, the significance of the stem *kum- is such that Christopher J. Beckwith mentions that Koreanic, Japonic and Old Chinese had the stem *kum- as a stem, denoting “a bear” (in case of the Old Chinese language, *kum- is only present as one of variants of stems, denoting “a bear”, while there is also a reconstructible Old Chinese word for “a bear”, which is related to the Tibeto-Burman word for “a bear”), but Tibeto-Burman does not share this stem *kum- for “a bear” with Old Chinese in spite of their relationship within Sino-Tibetan. In “The deep population history of northern East Asia from the Late Pleistocene to the Holocene”, the yDNA O1b2-L682 Japanese individual participated in the genetic cline, also involving yDNA N-M128 and yDNA O-F114>O-F16241 individuals, which may refer to a route from China to Japan for yDNA O-F114>O-F16241 and yDNA N-M128 individuals, bypassing the territories of the future Paekche Kingdom and Gaya(Kara) Confederacy in Southwestern Korea. Interactions along this geographic route, involving the yDNA O1b2-L682-related population of deep yDNA O1b2-L682* branches, one of whose descendants clustered with the Japanese on the PCA, but not with the Koreans, may explain the claimed appearance of the stem *kum- for “a bear”, shared by Japonic and Koreanic languages, in the Old Chinese language, while such a stem is absent in Tibeto-Burman languages, because yDNA O1b2-L682-related populations did not directly interact with the Tibeto-Burmans.

Christopher J. Beckwith reported some stems, which were allegedly shared between a Japonic-related language, the Old Chinese language and some Tibeto-Burman languages, implying that a Japonic-affiliated language, a few geographically southern Tibetan languages (e.g., the ancient Pyu language of Myanmar and the Old Tibetan language) and some form of the Old Chinese language might have even been in a form of a contact relationship. The most important cited word is a numeral, denoting “ten (10)”, for which Christopher J. Beckwith reported two forms for Proto-Japonic, and only one of these Proto-Japonic forms was similar to Old Chinese, Pyu and Old Tibetan forms. In “The deep population history of northern East Asia from the Late Pleistocene to the Holocene”, the yDNA O-F114>O-F16241 Japanese individual participated in the genetic cline, also involving a mentioned yDNA N-M128 individual and a mentioned yDNA O1b2-L682 Japanese individual. Since on the one hand, in “The deep population history of northern East Asia from the Late Pleistocene to the Holocene”, an yDNA O-M134>O-F114 individual clustered very close to yDNA N-M231-related individuals of the Shandong’s Houli culture, which, despite having features of round-bottomed fu cauldrons, similar to some features of round-bottomed fu cauldrons of the Hemudu culture of the Lower Yangtze River basin, had become quite geographically close to the homeland of the Sinitic ancestors of the Han Chinese in the Middle Yellow River, and, on the other hand, in “Human genetic history on the Tibetan Plateau in the past 5100 years”, the ancient ca. 4800-year-old yDNA O-M134>O-F114 individual appeared to be one of the most ancient Tibetan-related individuals to reach the elevation of the Qinghai-Tibetan Plateau, then the yDNA O-M134>O-F114 -related population should be one of Sinitic yDNA O-M134-related populations, providing a later link between the Sinitic part of the Yellow River basin and different Tibetan-related populations. Indeed, in “The deep population history of northern East Asia from the Late Pleistocene to the Holocene”, the yDNA O-F114>O-F16241 Japanese individual formed his own genetic cline of a very distant relationship with Tibetan-speaking Nepalese individuals, who in their turn participated in genetic clusters, leading on the PCA to specimens, belonging to speakers of various groups within Tibetan languages, and the mtDNA D4a4a of the mentioned yDNA O-F114>O-F16241 Japanese individual had a mutation C9569T, which appears in mtDNA M49, which is listed as one of Myanmar’s mtDNA lineages in “Ancient Mitogenomes Reveal the Origins and Genetic Structure of the Neolithic Shimao Population in Northern China”.

Consequently, if relevant common features with “global” cultures from China are omitted for simplicity, the position of the yDNA N-M128 Japanese individuals’ ancestors’ arrival to Japan should have been the following in “The deep population history of northern East Asia from the Late Pleistocene to the Holocene”: taking into account that their yDNA N-M128-related distant relative in China autosomally clustered with more southern individuals of more rice farming Jiangsu and Anhui Provinces due to neolithic circumstances, they had been arriving in accordance with the cline involving mtDNA D4a4a/yDNA O-F16241 Japanese individual, whose yDNA O-F17584 branch had a similar TMRCA (https://www.theytree.com/tree/O-F17584) to the TMRCA of yDNA N-M128, and who descended from yDNA O-F114-related neighbours of yDNA N-M231-related populations, and, taking into account that such researchers as Beckwith and some others claim the Proto-Japonic period for some of stems, shared by different mentioned language families, ancestors of both Japanese yDNA N-M128 individuals and Japanese yDNA O-F16241 individuals should have interacted with an expanding population, ancestral to ”Paekche/Gaya/Ara”-related yDNA O1b2-L682-related individuals (whose yDNA O1b2-L682/mtDNA F1a1b Japanese individual clustered with the main Japanese population) to explain the formation of the genetic cline of “The deep population history of northern East Asia from the Late Pleistocene to the Holocene”.

[AimSmall]

Scaled

Emperor_Wu:HRR1506796,0.027318,-0.405196,0.05506,-0.039729,-0.026774,-0.023427,0.013866,0.012923,-0.002045,0.013485,-0.055374,-0.004946,0.0055,0.002064,0,0.002917,0,-0.006841,0.002137,0.009379,-0.01148,-0.011252,-0.021815,-0.003133,0.004431





https://imgur.com/7OIJmZn

[ph2ter]

The similarity map also here:

[CLTVTE]

In its data, the article “Bronze and Iron Age population movements underlie Xinjiang population history” analyzes numerous cases of ancestry percentages, identical to ancestry percentages, which were discovered in “Northeastern Asian and Jomon-related genetic structure in the Three Kingdoms period of Gimhae, Korea” by Pere Gelabert, Choongwon Jeong and others, and which were observed, for example, in the series of ancient specimens “Korea-TK_2” (a male of yDNA D-M64 and a female of mtDNA B4c1a1a1a, buried in the Daeseong-dong tumulus of Gimhae, an area of the Gaya Confederacy). “The most important excavated center of the Gaya culture is a massive 3,700 m2 burial complex of rulers in Daeseong-dong in Gimhae, dated to the 1st–5th centuries CE.”

Interestingly, two models for Korea-TK_2, having the highest p-values, show that their p-values are rather similar, the first one being just slightly higher than the second one:

Korea-TK_2: [14,4% Japan_Jomon] [85,6% Japan_Kofun] p-value=0,898103
Korea-TK_2: [33,1% Japan_Jomon] [66,9% West_Liao_River_Bronze_Age (Upper Xiajiadian)] p-value=0,896472

The materials of the article “Bronze and Iron Age population movements underlie Xinjiang population history” allow to think that a very ancient common autosomal component, remaining in China, but even differently reaching areas outside China, evolved differently in the initially inland population, contributing to Japan_Kofun, and in the initially sea coastal population, contributing to West_Liao_River_Bronze_Age (the Upper Xiajiadian culture). However, since all non-peninsular continental male and female lineages in ancient Japan preserved closer relatives, who had remained in mainland China, especially including the area of the Yangtze River basin and areas of paths via areas of modern Jiangsu, Henan Provinces and finally Shandong, the Japan_Kofun as a source for one of the Gaya Confederacy’s population groups (Korea-TK_2) prevailed over the West_Liao_River_Bronze_Age (the Upper Xiajiadian culture) in those models.