https://www.biorxiv.org/content/10.1101/2020.03.25.004606v1 The Genomic Formation of Human Populations in East Asia
View ORCID ProfileChuan-Chao Wang, Hui-Yuan Yeh, Alexander N Popov, Hu-Qin Zhang, Hirofumi Matsumura, Kendra Sirak, Olivia Cheronet, Alexey Kovalev, Nadin Rohland, Alexander M Kim, Rebecca Bernardos, Dashtseveg Tumen, Jing Zhao, Yi-Chang Liu, Jiun-Yu Liu, Matthew Mah, Swapan Mallick, Ke Wang, Zhao Zhang, Nicole Adamski, Nasreen Broomandkhoshbacht, Kimberly Callan, Brendan J Culleton, Laurie Eccles, Ann Marie Lawson, Megan Michel, Jonas Oppenheimer, Kristin Stewardson, Shaoqing Wen, Shi Yan, Fatma Zalzala, Richard Chuang, Ching-Jung Huang, 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, Msato Nishino, Ken-ichi Shinoda, Olga A Shubina, Jianxin Guo, Qiongying Deng, Longli Kang, Dawei Li, Dongna Li, Rong Lin, Wangwei Cai, 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
doi: https://doi.org/10.1101/2020.03.25.004606
Abstract
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 191 individuals from Mongolia, northern China, Taiwan, the Amur River Basin and Japan dating to 6000 BCE - 1000 CE, many from contexts never previously analyzed with ancient DNA. We also report 383 present-day individuals from 46 groups mostly from the Tibetan Plateau and southern China. We document how 6000-3600 BCE people of Mongolia and the Amur River Basin were from populations that expanded over Northeast Asia, likely dispersing the ancestors of Mongolic and Tungusic languages. In a time transect of 89 Mongolians, we reveal how Yamnaya steppe pastoralist spread from the west by 3300-2900 BCE in association with the Afanasievo culture, although we also document a boy buried in an Afanasievo barrow with ancestry entirely from local Mongolian hunter-gatherers, representing a unique case of someone of entirely non-Yamnaya ancestry interred in this way. The second spread of Yamnaya-derived ancestry came via groups that harbored about a third of their ancestry from European farmers, which nearly completely displaced unmixed Yamnaya-related lineages in Mongolia in the second millennium BCE, but did not replace Afanasievo lineages in western China where Afanasievo ancestry persisted, plausibly acting as the source of the early-splitting Tocharian branch of Indo-European languages. Analyzing 20 Yellow River Basin farmers dating to ~3000 BCE, we document a population that was a plausible vector for the spread of Sino-Tibetan languages both to the Tibetan Plateau and to the central plain where they mixed with southern agriculturalists to form the ancestors of Han Chinese. We show that the individuals in a time transect of 52 ancient Taiwan individuals spanning at least 1400 BCE to 600 CE were consistent with being nearly direct descendants of Yangtze Valley first farmers who likely spread Austronesian, Tai-Kadai and Austroasiatic languages across Southeast and South Asia and mixing with the people they encountered, contributing to a four-fold reduction of genetic differentiation during the emergence of complex societies. We finally report data from Jomon hunter-gatherers from Japan who harbored one of the earliest splitting branches of East Eurasian variation, and show an affinity among Jomon, Amur River Basin, ancient Taiwan, and Austronesian-speakers, as expected for ancestry if they all had contributions from a Late Pleistocene coastal route migration to East Asia.
Глубокая популяционная история Восточной Азии остается малоизученной из-за отсутствия данных древней ДНК и скудной выборки современных людей. Мы сообщаем данные по всему геному от 191 человека из Монголии, северного Китая, Тайваня, бассейна реки Амур и Японии, датируемые 6000 г. до н.э. - 1000 г. н.э., многие из контекстов, которые ранее никогда не анализировались с помощью древней ДНК. Мы также сообщаем о 383 современных индивидуумах из 46 групп, в основном с Тибетского нагорья и южного Китая. Мы документируем, как в период 6000-3600 до н.э. жители Монголии и бассейна реки Амур были из населения, которое распространилось по Северо-Восточной Азии, вероятно, распространяя предков монгольских и тунгусских народов. Во временном разрезе 89 представителей древнего населения Монголии мы мы показываем, как к 3300-2900 гг. до н.э. ямные степные скотоводы распространились с запада в связи с Афанасьевской культурой, хотя мы также документируем мальчика, похороненного в Афанасьевском кургане с происхождением исключительно от местных монгольских охотников-собирателей , представляющий уникальный случай, когда кто-то совершенно не-ямного происхождения был похоронен в соответствии с ямным обрядом. Второе распространение ямного генетического компонента произошло через группы, которые унаследовали около трети своих генов от европейских земледельцев, и которые во втором тысячелетии до н.э. почти полностью вытеснили несмешанных потомков ямников в Монголии, но не вытеснили потомков афанасьевцев в западном Китае, где афанасьевский генетический компонент сохранился, возможно выступив источником рано отделившейся тохарской ветви индоевропейских языков. Анализируя 20 земледельцев бассейна реки Хуанхэ, датируемых примерно 3000 годом до н. э., мы фиксируем население, которое было вероятным вектором распространения сино-тибетских языков как на Тибетское плато, так и на центральную равнину, где они смешались с южными земледельцами, чтобы сформировать предков ханьцев(китайцев). Мы показываем, что индивидуумы во временном разрезе 52 древних тайваньцев, охватывающем, по крайней мере, 1400 г. до н.э. - 600 г. н.э., соответствовали почти прямым потомкам первых земледельцев долины Янцзы, которые, вероятно, распространили австронезийские, Тай-Кадайские и Австроазиатские языки по Юго-Восточной и Южной Азии и смешавшись с людьми, с которыми они столкнулись, поспособствовали четырехкратному снижению генетической дифференциации в период возникновения сложных обществ. Наконец, мы приводим данные дзёмонских охотников-собирателей из Японии, которые представляют одну из самых ранних отделившихся ветвей восточно-евразийской вариации, и показываем сходство между населением дзёмона, бассейном реки Амур, древним Тайванем и Австронезийцами, как и ожидалось в плане генетики, если бы все они имели вклад от позднеплейстоценовой миграции по прибрежному пути в Восточную Азию.
Online Table 1: Newly reported ancient individuals
Master ID Date: One of two formats. (Format 1) 95.4% CI calibrated radiocarbon age (Conventional Radiocarbon Age BP, Lab number) e.g. 5983-5747 calBCE (6980±50 BP, Beta-226472). (Format 2) Archaeological context date, e.g. 2500-1700 BCE Locality Region Y chromosome haplogroup (called only for males with 1240K data and only if >50000 autosomal SNPs hit) mtDNA haplogroup only called if >2x coverage
AH1-17 2904-2862 calBCE (4230±25 BP, XA-15378) Wuzhuangguoliang, Shaanxi, China China n/a (female) ..
18R21263 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
S91 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
S118 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
S94 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
18R21262 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
18R21264 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
S123 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
18R21266 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
18R21268 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
S96 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
S122 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (female) n/a (exome capture)
S120 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
S97 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
18R21265 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
18R21267 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
S95 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
AH1-7 3348-3090 calBCE (4490±30 BP, XA-15382) Wuzhuangguoliang, Shaanxi, China China n/a (female) ..
S121 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
S119 3400-2800 BCE Wuzhuangguoliang, Shaanxi, China China n/a (exome capture) n/a (exome capture)
I13882 2000-1500 BCE Rokutsu Shell Mound Japan n/a (female) N9b
I13883 2000-1500 BCE Rokutsu Shell Mound Japan D1b1d1a N9b2a
I13884 2000-1500 BCE Rokutsu Shell Mound Japan n/a (female) N9b
I13885 2000-1500 BCE Rokutsu Shell Mound Japan n/a (female) N9b
I13886 2000-1500 BCE Rokutsu Shell Mound Japan D1b1d1 N9b1
I13887 2000-1500 BCE Rokutsu Shell Mound Japan D1b1d1 N9b1
I6341 3500-3200 BCE Funadomari Japan D N9b1
I12973 1388-1134 calBCE (3020±25 BP, UCIAMS-226529, PSUG-5432) Bulgan, Khutag undur sum, Marzyn Khutul tomb 1 Mongolia Q1a2a1c D4j
I12958 1619-1458 calBCE (3265±30 BP, UCIAMS-226527, PSUG-5430) [use instead of less precise and inconsistent non-AMS-date of 1187-516 calBCE (2685±110 BP, Le-7213)] Khovd, Bulgan sum, Khukh uzuur, barrow 2 Mongolia n/a (female) G2a1
I6221 3000-2500 BCE Bayankhongor aimag, Erdenetsogt sum, Shatar chuluu kurgan 2 Mongolia J1 U5a1+@16192
I6222 3316-2918 calBCE (4415±31 BP, OxA-36222) Bayankhongor aimag, Erdenetsogt sum, Shatar chuluu kurgan 2 Mongolia R1b1a1a2a1a R
I13957 2858-2505 calBCE (4095±25 BP, PSUAMS-6942) [use instead of less precise but consistent non-AMS-date of 3011-2480 calBCE (4180±100 BP, Le-7219)] Bayan-Ulgiii aimag, Ulaankhus sum, Kurgak govi barrow 1, infant burial in main grave Mongolia C2b1a1 G2a+152
I13180 3013-2876 calBCE (4290±35 BP, PSUAMS-6944) [we used this direct AMS date on the same petrous bone analyzed for DNA instead of a previous and marginally non-overalpping AMS date of 2866-2577 calBCE (4114±29 BP, OxA-36230) on a different skeletal element supposed to be from the same individual, or a non-AMS date of 2572-2151 calBCE (3900±70 BP, Le-7212) also a different skeletal element supposed to be from the same individual] Bayan-Ulgii aimag, Ulaankhus sum, Kumdii govi barrow, burial 2 Mongolia n/a (female) A+152+16362
I12977 2913-2777 calBCE (4250±25 BP, UCIAMS-226551, PSUG-5466) [use instead of less precise and inconsistent non-AMS date of 2471-1781 calBCE (3725±115 BP, Le-7220)] Bayan-Ulgiii aimag, Ulaankhus sum, Kulala Ula barrow 1, secondary burial Mongolia C2b1a1 D4b1
I6361 2618-2487 calBCE (4034±16 BP, GrM-12938) [use instead of less precise and inconsistent non-AMS date of 2471-2044 calBCE (3825±70 BP, Le-7215)] Bayan-Ulgii aimag, Ulaankhus sum, Kurgak govi barrow 2 secondary burial Mongolia n/a (female) A+152+16362
I12978 2570-2476 calBCE [Rcombine of 2567-2468 calBCE (3983±17 BP, GrM-12984) and 2833-2487 calBCE (4050±25 BP, UCIAMS-226530, PSUG-5433); use instead of less precise and inconsistent non-AMS dates of 2461-2152 calBCE (3838±40 BP, Le-6932, Le-6933, Le-6939 R_Combine)] Khovd, Bulgan sum, Yagshiin hodoo barrow 1, bone find B Mongolia R1b1a1a2a2 C4+152
I12957 2571-2464 calBCE (3980±25 BP, UCIAMS-226526, PSUG-5429) Khovd, Bulgan sum, Yagshiin khodoo, Barrow 3 Mongolia R1b1a1a2a2 T1a1
I6369 752-416 calBCE (2455±20 BP, PSUAMS-3114) Khentii, Murun sum, Bus khairkhan Tomb 2 Mongolia n/a (female) B5b
I6353 1002-909 calBCE [Rcombine of 1002-846 calBCE (2780±25 BP, PSUAMS-2946), 1072-903 calBCE (2826±30 BP, OxA-36456)] Uvurkhangai, Khujirt sum, Shunkhlai Uul, Tomb 9 Mongolia Q1a1a1 G2b2
I12969 1119-980 calBCE (2875±20 BP, UCIAMS-226545, PSUG-5460) Khentii, Delgerkhaan sum, Khanan 3 Mongolia Q1a1a1 F1b1+@152
I13178 1042-916 calBCE (2825±20 BP, UCIAMS-226531, PSUG-5434) Khentii, Bayan-Adraga sum, Shuusyn boom Tomb 6 Mongolia n/a (female) R
I13958 1023-912 calBCE (2820±20 BP, UCIAMS-226558, PSUG-5473) Tuv aimag, Altanbulag sum, Erdenekhairkhan uulyn ar, Tomb 3 Mongolia n/a (<50000 SNPs) H2a
I7032 976-838 calBCE (2765±20 BP, PSUAMS-4004) Khentii, Batshireet sum, Tugsiin uvdug Tomb 1 Mongolia n/a (female) C4a2a1
I12971 976-838 calBCE (2765±20 BP, UCIAMS-226547, PSUG-5462) Dornod aimag, Gurvanzagal sum, Khadan khuv Tomb 1 Mongolia n/a (female) D4j7
I13963 968-833 calBCE (2750±20 BP, PSUAMS-6943) Khentii, Delgerkhaan sum, Rashaant tolgoi, tomb 6 Mongolia n/a (female) C5b1b
I6368 791-569 calBCE [Rcombine of 784-544 calBCE (2510±20 BP, PSUAMS-3907) and 800-571 calBCE (2550±20 BP, PSUAMS-4252)] Khentii, Delgerkhaan sum, Khanan Tomb 1a Mongolia n/a (female) L3
I6349 895-801 calBCE (2675±20 BP, PSUAMS-7398) Sükhbaatar aimag, Dariganga sum, Baaran-Khavtsal Mongolia Q1a1a1 D4m2
I6352 1071-925 calBCE (2845±20 BP, PSUAMS-7276) Uvurkhangai, Khujirt sum, Shunkhlai Uul, Grave #1 Mongolia Q1a1a F1b1b
I6365 814-783 calBCE (2610±20 BP, PSUAMS-7282) Khuvsgul aimag, Burentogtoh sum, Uushigiin uvur, slab grave 1 Mongolia N1c1a D4o1
I6359 1100-400 BCE [range based on other dates from the same cultural context and genetic cluster] Khuvsgul aimag, Alag-Erdene sum, Erhel nuur, Ulaan tolgoi Mongolia Q1a2a1c C5c
I6357 1100-400 BCE [This range is based on other dates from the same cultural context and genetic cluster. We ignore the direct date of 1277-1383 calCE (680±15 BP, PSUAMS-7278) on a post-cranial element for this individual because it mismatches the archaeological chronology as well as the genetic data (which match each other). The post-cranial was sampled from a box separate from the petrous bone analyzed for ancient DNA, which in the context of the mismatch with the genetic and archaeological context makes it plausible that the post-cranial element is not from the individual analyzed for DNA.] Tuv aimag, Altabulag sum, Tariat Brigad Tomb 5 Mongolia n/a (female) D4b2
I6364 1023-912 calBCE (2820±20 BP, PSUAMS-7281) Khovd, Bulgan sum, Uliastain dood denj I, kurgan 4 Mongolia n/a (female) M9a1a1c1
I13504 487-391 calBCE (2360±20 BP, UCIAMS-226556, PSUG-5471) Khuvsgul aimag, Ulaan-Uul sum Mongolia Q1a2a W
I6230 349-97 calBCE (2140±20 BP, PSUAMS-7396) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #47, Burial 2 Mongolia Q1a2a1c1 U2e1h
I7029 354-114 calBCE (2155±20 BP, PSUAMS-3877) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave 53, burial 2 Mongolia Q1a2a1c1 U4a1
I7026 356-177 calBCE (2180±15 BP, PSUAMS-4503) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #41, Burial 6 Mongolia n/a (<50000 SNPs) C4a1a+195
I6231 358-174 calBCE (2180±20 BP, PSUAMS-7397) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #47, Burial 3 Mongolia Q1a2a1c1 F1b1b
I6226 360-184 calBCE (2190±25 BP, PSUAMS-7270) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #35, Burial 2 Mongolia n/a (female) H91
I6224 361-203 calBCE (2210±20 BP, PSUAMS-7268) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #33, Burial 3 Mongolia R1a1a1b T2b34
I6233 361-203 calBCE (2210±20 BP, PSUAMS-7272) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery, Grave #49, Burial 1 Mongolia R1a1a1b2 F1b1+@152
I7023 365-204 calBCE (2215±20 BP, PSUAMS-4001) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #31, Burial 6 Mongolia n/a (female) H
I7028 381-208 calBCE (2235±20 BP, PSUAMS-4597) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave 46, burial 1 Mongolia n/a (female) H2a2a
I6232 387-209 calBCE (2245±20 BP, PSUAMS-7271) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #48, Burial 6 Mongolia Q1a2a1c1 U4b1a1a1
I7030 391-209 calBCE (2250±20 BP, PSUAMS-3722) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave 56 Mongolia R1a1a1b D4b1a2a1
I7022 391-209 calBCE (2250±20 BP, PSUAMS-4000) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #11, Burial 7 Mongolia Q1a1b1 C4a1a+195
I7024 391-209 calBCE (2250±20 BP, PSUAMS-4002) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #36, Burial 2 Mongolia R1a1a1b2 R1b1
I7027 399-235 calBCE (2275±20 BP, PSUAMS-4003) [used the PSUAMS date on the same skeletal element analyzed for ancient DNA instead of the non-overlapping measurement on an independent sample from the same individual of 358-195 calBCE (2188±15 BP, GrM-12941) published in Wilkins et al. Nature Ecology and Evolution 2020] Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #45, Burial 2 Mongolia R1a1a1b T1a1
I6223 400-200 BCE Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave 1 burial 5 Mongolia n/a (<50000 SNPs) ..
I6225 400-200 BCE [This range is based on other dates from the same cultural context and a brother from the same genetic cluster. We ignore the direct date of 1299-1404 calCE (605±20 BP, PSUAMS-7269) on a post-cranial element for this individual because it mismatches the archaeological chronology as well as the genetic data (which match each other). The post-cranial was sampled from a box separate from the petrous bone analyzed for ancient DNA, which in the context of the mismatch with the genetic and archaeological context makes it plausible that the post-cranial element is not from the individual analyzed for DNA.] Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #33, Burial 5 Mongolia R1a1a T2b34
I6356 400-200 BCE [This range is based on other dates from the same cultural context and genetic cluster. We ignore the direct date of 1386-1217 calBCE (3030±20 BP, PSUAMS-7277) on a mandible for this individual because it mismatches the archaeological chronology as well as the genetic data (which match each other). The mandible sampled for DNA was different from the petrous bone used for genetic analysis; these skeletal elements were from a commingled burial so it is plausible that the date does not correspond to the genetically analyzed individual.] Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery, Grave #58, Burial 2 Mongolia Q1a2a C4a1a3
I12970 400-235 calBCE (2280±20 BP, UCIAMS-226546, PSUG-5461) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave 48 burial 7 Mongolia Q1a2a1c1 A8a
I12956 405-364 calBCE (2305±20 BP, UCIAMS-226544, PSUG-5459) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave 48 burial 4 Mongolia n/a (<50000 SNPs) n/a (<2x)
I13965 397-233 calBCE (2270±20 BP, UCIAMS-226560, PSUG-5475) Bayan-Ulgiii aimag, Tsengel sum, Biluut 2 barrow 4 Mongolia n/a (female) G2a1
I6263 358-181 calBCE (2185±20 BP, PSUAMS-7273) Bayan Ulgii aimag, Tsengel sum, Tsagaan-Uul, Khuvsgol Mongolia n/a (female) H5a1
I6228 3-120 calCE (1950±20 BP, PSUAMS-7395) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery Grave #44 Mongolia C2b1a1b1 D4j12
I14194 971-834 calBCE (2755±20 BP, UCIAMS-226562, PSUG-5476) Sükhbaatar aimag, Tuvshinshiree sum, Ulaanzukh, Tomb 32 Mongolia n/a (female) C4a1
I13175 383-535 calCE (1625±20 BP, UCIAMS-226554, PSUG-5469) Khovd, Bulgan sum, Uliastai deed denj II kurgan 1 burial 1 (secondary) Mongolia R1b1c D4b1a2a
I13174 1500-1423 calBCE (3185±20 BP, UCIAMS-226564, PSUG-5483) [use instead of less precise but consistent non-AMS date of 1493-1091 calBCE (3050±70 BP, Le-6638)] Khovd, Munkhkhairkhan sum, Khukh tolgoi barrow 9 Mongolia Q1a2a1c U5a2
I13959 1441-1306 calBCE (3125±20 BP, UCIAMS-226559, PSUG-5474) [use instead of less precise but consistent non-AMS date of 1972-1306 calBCE (3340±130 BP, Le-6637)] Khovd, Munkhkhairkhan sum, Ulaan tolgoi, barrow 3 Mongolia n/a (female) n/a (<2x)
I12972 1447-1309 calBCE (3130±20 BP, UCIAMS-226553, PSUG-5468) Sükhbaatar aimag, Tuvshinshiree sum, Adgiin gol Tomb 21 Mongolia Q1a1 D4i
I14037 1420-1296 calBCE (3095±20 BP, UCIAMS-226561, PSUG-5475) Sükhbaatar aimag, Tuvshinshiree sum, Bulgiin ekh, Tomb 47 Mongolia Q1a1a M11b
I13964 1500-800 BCE Sükhbaatar aimag, Tuvshinshiree sum, Ulaanzukh, Tomb 17 Mongolia n/a (female) C4a1a+195
I12960 1437-1304 calBCE (3120±20 BP, UCIAMS-226528, PSUG-5431) Uvurkhangai, Khujirt sum, Shunkhlai Uul, Tomb 2 Mongolia Q1a1 C4a1a+195
I12506 1438-1299 calBCE (3115±25 BP, PSUAMS-6933) [use instead of previous date of 1608-1416 calBCE (3215±40 BP, OxA-36459) not from the same skeletal element] Sükhbaatar aimag, Tuvshinshiree sum, Ulaanzukh, Tomb 42 Mongolia n/a (<50000 SNPs) n/a (<2x)
I12976 1437-1304 calBCE (3120±20 BP, UCIAMS-226550, PSUG-5465) [use instead of less precise but consistent non-AMS date of 1396-845 calBCE (2910±110 BP, Le-7226)] Bayan-Ulgiii aimag, Ulaankhus sum, Kulala Ula barrow 2 Mongolia N1 U2e2a1a2
I7033 1209-1038 calBCE (2920±20 BP, PSUAMS-4005) Uvs aimag, Ulaangom sum, Chandman Uul, Ulaangom cemetery kurgan 59 Mongolia R1a1a1b R
I6363 1214-1058 calBCE [Rcombine of 1210-1047 calBCE (2925±20 BP, PSUAMS-7280) and 1277-1057 calBCE (2967±31 BP, OxA-X-2737-54); do not use the less precise but consistent non-AMS date of 1208-841 calBCE (2840±70 BP, Le-7979)] Khovd, Bulgan sum, Khudzrtyn gol II, barrow 1, burial 4 (main burial) Mongolia R1a1a1b2a2b U2e1
I6351 1003-854 calBCE (2785±20 BP, PSUAMS-7275) Uvs aimag, Khyargas sum, Urgun shireg, Tomb 3 Mongolia n/a (female) C4b1
I13505 1119-980 calBCE (2875±20 BP, UCIAMS-226557, PSUG-5472) Bayankhongor aimag, Erdentsogt sum Mongolia Q1a1a1 A+152+16362
I7039 1209-1038 calBCE (2920±20 BP, PSUAMS-4006) Khovd, Duut sum, Khoit Tsenkheriin gol 1988 barrow 3 Mongolia Q1a2a1c D4j
I13768 1210-1047 calBCE (2925±20 BP, PSUAMS-7115) Bayankhongor aimag, Ulziit sum, Muunit uul Mongolia N D4j1
I12975 1226-1056 calBCE (2950±20 BP, UCIAMS-226549, PSUG-5464) Khuvsgul aimag, Khankh sum, Khavtsal II Tomb 1 (Khirgisuur) Mongolia C2b1a U5a1a2a
I13767 1374-1131 calBCE (3005±20 BP, UCIAMS-226534, PSUG-5437) Bayankhongor aimag, Ulziit sum, Khar Sair am Mongolia Q1a2a1c R1b1
I13766 1381-1131 calBCE (3010±25 BP, UCIAMS-226533, PSUG-5436) Bayankhongor aimag, Erdentsogt sum Mongolia Q1a2a1c C5b1b
I6262 1396-1231 calBCE (3050±20 BP, PSUAMS-3111) Bayankhongor aimag, Khar Sairiin Am, Khirigsuur Mongolia n/a (female) A8a1
I6264 1397-1236 calBCE (3055±20 BP, PSUAMS-3112) Bayankhongor aimag, Khar Sairiin Am, Khirigsuur Mongolia n/a (female) A8a1
I6347 1411-1284 calBCE (3080±20 BP, PSUAMS-3113) Uvs aimag, Züünkhangai sum, Savaa Khar Mongolia n/a (female) K2a5
I6362 1015-911 calBCE (2815±20 BP, PSUAMS-2955) Khovd, Bulgan sum, Khudzhirtyn gol I, kurgan 2 Mongolia n/a (female) D4b1
I6367 1258-1088 calBCE (2955±20 BP, PSUAMS-7284) [use instead of less precise but agreeing date of 1260-1040 calBCE (2820±30 BP, Beta-306037)] Bayan-Ulgii aimag, Tsengel sum, Biluut 2, barrow 3 Mongolia O2a2b1a1a C4a1
I12955 1948-1777 calBCE (3545±20 BP, UCIAMS-226543, PSUG-5458) Zavkhan, Bayantes sum, Khukh Khushoony bom, Barrow 3 Mongolia N1c1a D4i
I6348 1746-1630 calBCE (3395±25 BP, PSUAMS-7274) Bulgan, Khutag-Undur sum, Haruulyn Gozgor tomb 1-113 Mongolia Q1a2a1c HV12b1
I13173 1879-1694 calBCE (3460±20 BP, UCIAMS-226552, PSUG-5467) [use instead of less precise and inconsistent non-AMS date of 1609-1232 calBCE (3150±70 BP, Le-6936)] Khovd, Munkhkhairkhan sum, Ulan goviin uzuur barrow 2 Mongolia N H15b1
I13176 890-982 calCE (1115±20 BP, UCIAMS-226555, PSUG-5470) Khovd, Bulgan sum, Hudzhirtyn gol II, kurgan 1 burial 3 (secondary) Mongolia R1b1c D4i
I12974 1225-1285 calCE (750±20 BP, UCIAMS-226548, PSUG-5463) Khentii, Murun sum, Bus khairkhan Tomb 3 Mongolia n/a (female) C4a1a+195
I13961 1282-1388 calCE (660±15 BP, PSUAMS-7114) Dornod aimag, Gurvanzagal sum Tsagaan chuluut kurgan 1 Mongolia n/a (female) A+152+16362
I7021 5211-5000 calBCE (6145±35 BP, PSUAMS-3931) Dornod aimag, Bulgan sum, Norovlin Uul burial Mongolia C2b1a1 D4h4
I7031 4458-4364 calBCE (5590±20 BP, PSUAMS-4598) Dornod aimag, Bayantumen sum, Kherlengiin baruun ereg (Kerulen River right bank), Grave 2 (1968 y.) Mongolia n/a (<50000 SNPs) C5b1
I6358 5974-5747 calBCE (6965±35 BP, PSUAMS-7279) Dornod aimag, Bayantumen sum, Kherlen River South bank nearby Choibalsan town Mongolia n/a (female) F1b1+@152
I13698 5700-5000 BCE Bulgan, Khutag-Undur sum, Egiin-gol river valley, Marzyn Khutul, tomb 1-055-1, sceleton 2 Mongolia C2b1a1 F1b1+@152
I13701 5721-5614 calBCE (6739±40 BP, AA-107617) Bulgan, Khutag-Undur sum, Egiin-gol river valley, Kharuulyn gozgor, tomb 1-116 Mongolia n/a (female) C4a1a3
I13179 5607-5477 calBCE (6565±30 BP, UCIAMS-226532, PSUG-5435) Bulgan, Khutag undur sum, Kharuulyn gozgor, tomb 1-116 Mongolia n/a (female) C4a1a3
I14000 5575-5474 calBCE (6571±47 BP, COL3897.1.1) Bulgan, Khutag-Undur sum, Egiin-gol river valley, Kharuulyn gozgor, tomb 1-117-1, sceleton 3 Mongolia n/a (female) D4j
I11696 5632-5531 calBCE (6655±30 BP, UCIAMS-226523, PSUG-5426) Bulgan, Khutag-Undur sum, Marzyn khutul, Tomb 1-055-1, sceleton 1 Mongolia C2b1a1 D4j
I11698 5632-5515 calBCE (6645±35 BP, UCIAMS-226525, PSUG-5428) Bulgan, Khutag-Undur sum, Marzyn khutul, Tomb 1-055-1, sceleton 2 Mongolia C2b1a1 D4b1c
I11697 5617-5491 calBCE (6610±30 BP, UCIAMS-226524, PSUG-5427) Bulgan, Khutag-Undur sum, Marzyn khutul, Tomb 1-055-1, sceleton 4 Mongolia C2b1a1 D4j8
I6366 1410-1261 calBCE (3065±25 BP, PSUAMS-7283) Bayan-Ulgiii aimag, Tsengel sum, Biluut 2 barrow 2 Mongolia n/a (female) J2b1a2a
I1208 450-750 CE Primorsky Krai, Chernyatino 5, Mohe-Bohai Graveyard Russia n/a (<50000 SNPs) H2
I14819 3750-3600 BCE [layer date based on mothering children with I3356 at 3696-3638 calBCE (4875±20 BP, PSUAMS-2160)] Primorsky Krai, Boisman-2 Russia n/a (female) C5a
I14308 5400-3600 BCE Primorsky Krai, Boisman-2 Russia C2a1a2(xC2a1a2a) D4o2a
I3356 3696-3638 calBCE (4875±20 BP, PSUAMS-2160) Primorsky Krai, Boisman-2 Russia C2a1a3(xC2a1a3a) C5b1
I3354 5371-5219 calBCE (6325±35 BP, PSUAMS-2070) Primorsky Krai, Boisman-2 Russia n/a (female) D4
I1196 5213-5008 calBCE (6155±35 BP, Poz-83476) Primorsky Krai, Boisman-2 Russia n/a (female) C5b1
I1206 5213-4834 calBCE (6080±70 BP, TERRA-b06129705) Primorsky Krai, Boisman-2 Russia n/a (female) R
I1198 5034-4806 calBCE (6030±40 BP, Poz-83478) Primorsky Krai, Boisman-2 Russia n/a (female) D4l2
I1193 4981-4795 calBCE (5995±35 BP, Poz-83474) Primorsky Krai, Boisman-2 Russia C2a1a3 C5b
I3355 4981-4787 calBCE (5985±35 BP, PSUAMS-2114) Primorsky Krai, Boisman-2 Russia C2a1a(xC2a1a1,C2a1a2,C2a1a3) C5a
I1197 4956-4729 calBCE (5970±40 BP, Poz-83477) Primorsky Krai, Boisman-2 Russia n/a (female) C5a
I1194 4792-4604 calBCE (5840±35 BP, Poz-83475) Primorsky Krai, Boisman-2 Russia n/a (female) C5a
I1190 4721-4335 calBCE (5860±100 BP, TERRA-06129717) Primorsky Krai, Boisman-2 Russia n/a (female) D4e5
I1192 4932-4608 calBCE (5890±60 BP, TERRA-b06129733) Primorsky Krai, Boisman-2 Russia C2a1(xC2a1a1a,C2a1a2a) B4d1'2'3
I14307 5400-3600 BCE Primorsky Krai, Boisman-2 Russia n/a (female) D4o2a
I14772 3750-3600 BCE [layer date based on being daughter of I3356 at 3696-3638 calBCE (4875±20 BP, PSUAMS-2160)] Primorsky Krai, Boisman-2 Russia n/a (female) C5a
I14774 3750-3600 BCE [layer date based on being daughter of I3356 at 3696-3638 calBCE (4875±20 BP, PSUAMS-2160)] Primorsky Krai, Boisman-2 Russia n/a (female) C5a
I14773 3750-3600 BCE [layer date based on being son of I3356 at 3696-3638 calBCE (4875±20 BP, PSUAMS-2160)] Primorsky Krai, Boisman-2 Russia n/a (<50000 SNPs) C5a
I14771 3750-3600 BCE [layer date based on being son of I3356 at 3696-3638 calBCE (4875±20 BP, PSUAMS-2160)] Primorsky Krai, Boisman-2 Russia C2a(xC2a1a1,C2a1a2a) C5a
I1209 1039-1224 calCE (880±35 BP, Poz-83439) Roshino-4, Primorsky Krai Russia O2 D4a1e
I3358 1050-1220 calCE (875±20 BP, PSUAMS-2115) Roshino-4, Primorsky Krai Russia n/a (female) D4a1
I15926 1800-1950 CE Nevelsk 2 Russia D1a2b(xD1a2b1) N9b1
I1202 996-830 calBCE (2760±35 BP, Poz-83438) Pospelovo 1, Primorsky Krai Russia N1a(xN1a1,N1a2) C5b
I13721 250-650 CE Green Island, Gongguan site Taiwan n/a (female) Y2a1
I13722 250-650 CE Green Island, Gongguan site Taiwan n/a (<50000 SNPs) Y2a1
I3618 1402-1262 calBCE (3060±20 BP, PSUAMS-2268) Yilan, Hanben Taiwan O2a2b2a2 R
I8071 150-465 CE Yilan, Hanben Taiwan n/a (female) R
I13695 150-465 CE Yilan, Hanben Taiwan n/a (<50000 SNPs) F4b
I8076 150-465 CE Yilan, Hanben Taiwan n/a (<50000 SNPs) R30
I8080 150-465 CE Yilan, Hanben Taiwan O1a1a1a F3b1a+16093
I8075 150-465 CE Yilan, Hanben Taiwan n/a (female) E2b
I3611 150-465 CE Yilan, Hanben Taiwan n/a (female) E1a1a1
I3616 150-465 CE Yilan, Hanben Taiwan n/a (female) B4b1a2
I3615 150-465 CE Yilan, Hanben Taiwan n/a (female) R
I3620 150-465 CE Yilan, Hanben Taiwan n/a (female) R
I3617 150-465 CE Yilan, Hanben Taiwan n/a (female) F4b1
I3621 150-465 CE Yilan, Hanben Taiwan n/a (female) R9c1b2
I15159 150-465 CE Yilan, Hanben Taiwan n/a (female) F4b1
I3614 150-465 CE Yilan, Hanben Taiwan O2a2b2a R
I3612 150-465 CE Yilan, Hanben Taiwan O2a2b2a2 E1a1a1
I3613 150-465 CE Yilan, Hanben Taiwan O1a1a1a1 R
I3619 395-536 calCE (1610±25 BP, PSUAMS-2075) Yilan, Hanben Taiwan n/a (female) F3b1a+16093
I14925 510-870 CE Yilan, Hanben Taiwan n/a (female) M7b1a2a1
I14934 510-870 CE Yilan, Hanben Taiwan O1a1a1a1 E2a
I14933 510-870 CE Yilan, Hanben Taiwan O1a2 F3b1a
I14931 510-870 CE Yilan, Hanben Taiwan O2a2 E1a1a1
I8072 510-870 CE Yilan, HanBen Taiwan O1a E1a1a
I14929 510-870 CE Yilan, Hanben Taiwan O2a2b B4a1a
I8074 510-870 CE Yilan, Hanben Taiwan n/a (female) F3b1a+16093
I8081 510-870 CE Yilan, Hanben Taiwan O1a1a1a F3b1a2
I15158 510-870 CE Yilan, Hanben Taiwan O1a2 B4b1a2f
I13692 510-870 CE Yilan, Hanben Taiwan n/a (female) M7c1c3
I3728 510-870 CE Yilan, Hanben Taiwan n/a (female) F3b1a+16093
I3736 510-870 CE Yilan, Hanben Taiwan O1a1a1a1 F4b1
I3732 510-870 CE Yilan, Hanben Taiwan n/a (female) F1a3a
I3735 510-870 CE Yilan, Hanben Taiwan n/a (female) E1a
I3727 510-870 CE Yilan, Hanben Taiwan n/a (female) D6a2
I14937 510-870 CE Yilan, Hanben Taiwan n/a (female) B4a1a
I3734 510-870 CE Yilan, Hanben Taiwan O2a2a1a2 E1a
I3731 510-870 CE Yilan, Hanben Taiwan O1a1a1a1a1 F3b1
I3733 510-870 CE Yilan, Hanben Taiwan O1a1a1a1 F3b1
I8078 510-870 CE Yilan, Hanben Taiwan O2a2b2 E1a1
I15156 510-870 CE Yilan, Hanben Taiwan n/a (<50000 SNPs) D6a2
I13697 150-465 CE Yilan, Hanben Taiwan n/a (<50000 SNPs) R
I15164 150-465 CE Yilan, Hanben Taiwan n/a (female) E2a
I15160 150-465 CE Yilan, Hanben Taiwan n/a (<50000 SNPs) F4b1
I15162 150-465 CE Yilan, Hanben Taiwan n/a (<50000 SNPs) E1a1a
I15166 150-465 CE Yilan, Hanben Taiwan n/a (female) E1a1a
I8079 150-465 CE Yilan, Hanben Taiwan n/a (female) F3b1
I14930 510-870 CE Yilan, Hanben Taiwan n/a (female) E1a1a1
I14928 510-870 CE Yilan, Hanben Taiwan n/a (female) B5a2a1+16129
I14926 510-870 CE Yilan, Hanben Taiwan n/a (female) F4b1
I13694 510-870 CE Yilan, Hanben Taiwan n/a (female) B4a1a
I14927 510-870 CE Yilan, Hanben Taiwan n/a (female) M7b1a2a1
I13696 510-870 CE Yilan, Hanben Taiwan n/a (<50000 SNPs) n/a (<2x)