Genetic history of East Asians

Source: Wikipedia, the free encyclopedia.

This article summarizes the genetic makeup and population history of

East Eurasians
" in population genomics.

Overview

Phylogenetic position of East Asian lineages among other Eastern Eurasians
Schematic of Populations in Eurasia from 45 to 10 kaBP
Highlighted regions show where ancient individuals associated with the labeled ancestry have been sampled.

Population genomic studies have studied the origin and formation of modern East Asians. Ancestors of East Asians (Ancient East Eurasians) split from other human populations possibly as early as 70,000 to 50,000 years ago. Possible routes into East Asia include a northern route model from Central Asia, beginning north of the Himalayas, and a southern route model, beginning south of the Himalayas and moving through Southeast Asia.[1][2]

Phylogenetic data suggests that an early

Papuans may have also received some geneflow from an earlier group (xOoA),[4] around 2%,[5] next to additional archaic admixture in the Sahul
region.

The southern route model for East Asians has been corroborated in multiple recent studies, showing that most of the ancestry of Eastern Asians arrived from the southern route in to Southeast Asia at a very early period, starting perhaps as early as 70,000 years ago, and dispersed northward across Eastern Asia.

Q and R, as well as Ancient North Eurasian ancestry.[12][13]

The southern migration wave likely diversified after settling within East Asia, while the northern wave, which probably arrived from the Eurasian steppe, mixed with the southern wave, probably in Siberia.[14]

A review paper by Melinda A. Yang (in 2022) described the 'East- and Southeast Asian' lineage (ESEA); which is ancestral to modern

Northern China, which can already be differentiated from the deeply related Ancestral Ancient South Indians (AASI) and Australasian (AA) lineages.[3]
There are currently eight detected, closely related, sub-ancestries in the ESEA lineage:

Proposed migration routes (Wang 2013) of East Asian paternal haplogroups (C, D, N, and O), during the peopling of East Asia[15]

Modern Northeast Asians derive most of their ancestry from the "Amur" (Ancient Northeast Asian) sub-linesge, which expanded massively with

rice cultivation. Contemporary East Asians (most notably Sino-Tibetan speakers) mostly have Yellow River ancestry, associated with both millet and rice cultivation. "East Asian Highlanders" (Tibetans) carry both Tibetan ancestry and Yellow River ancestry. Japanese people were found to have a tripartite origin; consisting of Jōmon ancestry, Amur ancestry, and Yellow River ancestry. Indigenous peoples of the Americas formed from Ancient North Eurasians and from an early Northern East Asian branch, giving rise to "Ancient Paleo-Siberians", which in turn gave rise to both "modern Paleosiberians" and contemporary Native Americans. Isolated hunter-gatherers in Southeast Asia, specifically in Malaysia and Thailand, such as the Semang, derive most of their ancestry from the Hoabinhian lineage.[16][17][18][19][20][21][22]

Proposed migration routes of maternal haplogroups during the peopling of Eurasia[23]

The genetic makeup of East Asians is primarily characerized by the Ancient Northern East Asian (ANEA) and Ancient Southern East Asian (ASEA) lineages, which diverged from each other at least 19,000 years ago, after the divergence of the Jōmon, Longlin, Hoabinhian and Tianyuan lineages.[24][3]

Ancient and historical populations

Xiongnu people

Main article: Xiongnu

The

Eurasian Steppe from the 3rd century BC to the late 1st century AD. Chinese sources report that Modu Chanyu, the supreme leader after 209 BC, founded the Xiongnu Empire.[26]

Autosomal DNA

It was found that the "predominant part of the Xiongnu population is likely to have spoken Turkic". However, important cultural, technological and political elements may have been transmitted by Eastern

Slab Grave culture.[30]

Paternal lineages

A review of the available research has shown that, as a whole, 53% of Xiongnu paternal haplogroups were East Eurasian, while 47% were West Eurasian.

haplogroup R1.[33][34]

Maternal lineages

The bulk of the genetics research indicates that, as a whole, 73% of Xiongnu maternal haplogroups were East Eurasian, while 27% were West Eurasian.[35] A 2003 study found that 89% of Xiongnu maternal lineages from the Egiin Gol valley were of East Asian origin, while 11% were of West Eurasian origin.[36] A 2016 study of Xiongnu from central Mongolia found a considerably higher frequency of West Eurasian maternal lineages, at 37.5%.[37]

Xianbei people

Main article: Xianbei

Autosomal DNA

A full genome study on multiple Xianbei remains found them to be derived primarily to exclusively from the Ancient Northeast Asian gene pool.[38]

Paternal lineages

A genetic study published in the American Journal of Physical Anthropology in August 2018 noted that the paternal

Rouran, and was probably an important lineage among the Donghu people.[39]

Maternal lineages

Genetic studies published in 2006 and 2015 revealed that the

East Asian origin. According to Zhou (2006) the maternal haplogroup frequencies of the Tuoba Xianbei were 43.75% haplogroup D, 31.25% haplogroup C, 12.5% haplogroup B, 6.25% haplogroup A and 6.25% "other".[40] Zhou (2014) obtained mitochondrial DNA analysis from 17 Tuoba Xianbei, which indicated that these specimens were, similarly, completely East Asian in their maternal origins, belonging to haplogroups D, C, B, A, O and haplogroup G.[41][42]

Jōmon people

Main article: Jōmon people

The Jōmon people represent the indigenous population of the Japanese archipelago during the Jōmon period. They are inferred to descend from the Paleolithic inhabitants of Japan. Genetic analyses on Jōmon remains found them to represent a deeply diverged East Asian lineage. The Jōmon lineage is inferred to have diverged from Ancient East Asians before the divergence between Ancient Northern East Asians and Ancient Southern East Asians, but after the divergence of the basal Tianyuan man and or Hoabinhians. Beyond their broad affinity with Eastern Asian lineages, the Jōmon also display a weak affinity for Ancient North Eurasians (ANE), which may be associated with the introduction of microblade technology to Northeast Asia and northern East Asia during the Last Glacial Maximum via the ANE or Ancient Paleo-Siberians.[43][44]

Hoabinhians

Main article:
Southern China. While the Upper Paleolithic origins of this 'Hoabinhian ancestry' are unknown, Hoabinhian ancestry has been found to be related to the main 'East Asian' ancestry component found in most modern East and Southeast Asians, although deeply diverged from it.[45][46] Together with the Paleolithic Tianyuan man, they form early branches of East Asian genetic diversity, and are described as "Basal Asian" (BA) or "Basal East Asian" (BEA).[47]

Modern populations

East Asian peoples

Manchu and Daur peoples

Main article: Tungusic peoples

Autosomal DNA

A study on the Manchu population of Liaoning reported that they have a close genetic relationship and significant admixture signals from northern Han Chinese. The Liaoning Manchu were formed from a major ancestral component related to Yellow River farmers and a minor ancestral component linked to ancient populations from the Amur River Basin, or others. The Manchu were therefore an exception to the coherent genetic structure of Tungusic-speaking populations, likely due to the large-scale population migrations and genetic admixtures in the past few hundred years.[49]

Paternal lineages

A plurality of

Aisin Gioro and is found in ten different ethnic minorities in northern China, but is largely absent from Han Chinese.[55][56][54] The Manchu people also display a significant amount of haplogroup C-M217, but the most often observed Y-DNA haplogroup among present-day Manchus is Haplogroup O-M122, which they share in common with the general population of China.[57][50][58]

Ainu people

Main article: Ainu people

The exact origins of the early Ainu remains unclear, but it is generally agreed to be linked to the

Epi-Jōmon period, with later influences from the nearby Okhotsk culture.[59] The Ainu appear genetically most closely related to the Jōmon period peoples of Japan. The genetic makeup of the Ainu represents a "deep branch of East Asian diversity". Compared to contemporary East Asian populations, the Ainu share "a closer genetic relationship with northeast Siberians".[6][44]

Japanese people

A population genomic PCA graph, showing the substructure of Eastern Asian populations
Main articles: Japanese people and Genetic and anthropometric studies on Japanese people

Japanese populations in modern Japan can be traced to three separate, but related demographics: the Ainu, Ryukyuan and Mainland Japanese (Yamato people). The populations are closely related to clusters found in North-Eastern Asia[60][61][62] with the Ainu group being most similar to Ryukyuans[61][63] and the Yamato group being most similar to Koreans[64][65][66] among other East Asian people.

Autosomal DNA

The majority of Japanese genetic ancestry is derived from sources related to other mainland Asian groups, mostly Koreans, while the other amount is derived from the local Jōmon hunter-gatherers.[67]

Geomic transitions in parallel with cultural transitions in pre- and protohistoric Japan

According to a full genome analyses, the modern Japanese harbor a Northeast Asian, an East Asian, and an indigenous Jōmon component. In addition to the indigenous Jōmon hunter-gatherers and the Yayoi period migrants, a new strand was hypothesized to have been introduced during the Yayoi-Kofun transition period that had strong cultural and political affinity with Korea and China.[68] Modern Japanese have inherited European ancestry, around 2.2%.[69]

Paternal lineages

A comprehensive study of worldwide Y-DNA diversity (Underhill et al. 2000) included a sample of 23 males from Japan, of whom 35% belonged to haplogroup D-M174, 26% belonged to O-M175, 22% belonged to O-M122, 13% belonged to C-M8 and C-M130, and 4.3% belonged to N-M128.[70] Poznik et al. (2016) reported the haplogroups of a sample of Japanese men from Tokyo:[71] 36% belonged to D2-M179, 32% had O2b-M176, 18% carried O3-M122, 7.1% carried C1a1-M8, 3.6% belonged to O2a-K18, and 3.6% carried C2-M217.[72]

Maternal lineages

According to an analysis of the

Z (3.4%), M9 (2.5%), and M8 (1.7%).[73]

Korean people

Main article: Koreans
Regional reference panel, PCA, and Admixture analysis

Modern Koreans are overall more similar to northeast Asians than to southeast Asians.[74] The reference population for Koreans used in Geno 2.0 Next Generation is 94% Eastern Asia and 5% Southeast Asia & Oceania.[75]

Autosomal DNA

Ancient genome comparisons revealed that the genetic makeup of Koreans can be best described as an admixture between Northern East Asian hunter-gatherers and an influx of rice-farming agriculturalists from the Yangtze river valley.[74] This is supported by archeological, historical and linguistic evidence, which suggest that the direct ancestors of Koreans were proto-Koreans who inhabited the northeastern region of China and the Korean Peninsula during the Neolithic (8,000–1,000 BC) and Bronze (1,500–400 BC) Ages.[76]

There is evidence for considerable genetic diversity,[77] including elevated levels of Jōmon ancestry among early southern Koreans.[78] It was hypothesized that the Jōmon ancestry of ancient Koreans was lost over time, as they continually mixed with incoming populations from northern China,[79] followed by a period of isolation during the Three Kingdoms period, resulting in the homogenous gene pool of modern Koreans.[80][77]

qpAdm analysis of ancient Koreans with Jōmon genes (Gelabert 2022)

A 2022 study was unable to detect significant Jōmon ancestry in modern Koreans, however by using different proxies of ancestry, a Jōmon contribution of 3.1–4.4% was found for present-day Ulsan Koreans. Nevertheless, the authors suggested that the model that yielded this result is not the most reliable.[81]

Evidence for both Southern and Northern mtDNA and Y-DNA haplogroups has been observed in Koreans, similar to Japanese.[48]

Paternal lineages

Studies of polymorphisms in the human Y-chromosome have so far produced evidence to suggest that the Korean people have a long history as a distinct, mostly endogamous ethnic group, with successive waves of people moving to the peninsula and three major Y-chromosome haplogroups.[82] A majority of Koreans belong to subclades of haplogroup O-M175 (ca. 79% in total,[76][83] with about 42%[83] to 44%[76] belonging to haplogroup O2-M122, about 31%[76] to 32%[83] belonging to haplogroup O1b2-M176, and about 2%[76] to 3%[83] belonging to haplogroup O1a-M119), while a significant minority belong to subclades of haplogroup C2-M217 (ca. 12%[76] to 13%[83] in total). Other Y-DNA haplogroups, including haplogroup N-M231, haplogroup D-M55, and haplogroup Q-M242, are also found in smaller proportions of present-day Koreans.[84][85][86]

Maternal lineages

Studies of Korean

Y, F, D5, M7, M8, M9, M10, M11, R11, C, and Z.[87][88][89]

Mongolic peoples

Main article: Mongolic peoples

The ethnogenesis of Mongolic peoples is largely linked with the expansion of

Onnigud. The Daur people are descendants of the para-Mongolic Khitan people.[92]

Paternal lineages

The majority of Mongols in Mongolia and Russia belong to subclades of haplogroup C-M217,[93] followed by lower frequency of O-M175 and N-M231.[94] A minority belongs to haplogroup Q-M242, and a variety of West Eurasian haplogroups.[95]

Maternal lineages

The maternal haplogroups are diverse but similar to other northern Asian populations, including Haplogroup D, Haplogroup C, Haplogroup B, and Haplogroup A, which are shared among indigenous American and Asian populations.[96]

Han Chinese

See also: Han Chinese

Han Chinese descend primarily from Neolithic Yellow River farmers, which formed primarily from Ancient Northern East Asians with some contributions from Ancient Southern East Asians. Northern Han Chinese mostly carry ANEA ancestry with a moderate degree of ASEA admixture, whereas southern Han Chinese carry significantly higher levels of ASEA ancestry than Northern Han, although ANEA ancestry still predominates.[97][24][98]

Autosomal DNA

A 2018 study calculated pairwise FST (a measure of genetic difference) based on genome-wide SNPs, among the Han Chinese (Northern Han from Beijing and Southern Han from Hunan, Jiangsu and Fujian provinces), Japanese and Korean populations sampled. It found that the smallest FST value was between Northern Han Chinese (Beijing) (CHB) and Southern Han (Hunan, Fujian, etc.) Chinese (CHS) (FST[CHB-CHS] = 0.0014), while CHB and Korean (KOR) (FST[CHB-KOR] = 0.0026) and between KOR and Japanese (JPT) (FST[JPT-KOR] = 0.0033). Generally, pairwise FST between Han Chinese, Japanese and Korean (0.0026~ 0.0090) are greater than that within Han Chinese (0.0014). These results suggested Han Chinese, Japanese and Korean are different in terms of genetic make-up, and the differences among the three groups are much larger than that between northern and southern Han Chinese.[99] Nonetheless, there is also genetic diversity among the Southern Han Chinese. The genetic composition of the Han population in Fujian might not accurately represent that of the Han population in Guangdong.

A PCA graph illustrates the genetic differences among Han Chinese groups.[100]

Another study shows that the northern and southern Han Chinese are genetically close to each other and it finds that the genetic characteristics of present-day northern Han Chinese were already formed prior to three thousand years ago in the Central Plain area.[101]

A recent genetic study on the remains of people (~4,000 years

BP) from the Mogou site in the Gansu-Qinghai (or Ganqing) region of China revealed more information on the genetic contributions of these ancient Di-Qiang people to the ancestors of the Northern Han. It was deduced that 3,300 to 3,800 years ago some Mogou people had merged into the ancestral Han population, resulting in the Mogou people being similar to some northern Han in sharing up to ~33% paternal (O3a) and ~70% maternal (D, A, F, M10) haplogroups. The mixing ratio was possibly 13–18%.[102]

The estimated contribution of northern Han to southern Han is substantial in both paternal and maternal lineages and a geographic cline exists for mtDNA. As a result, the northern Han are one of the primary contributors to the gene pool of the southern Han. However, it is noteworthy that the expansion process was not only dominated by males, as is shown by both contribution of the Y-chromosome and the mtDNA from northern Han to southern Han. Northern Han Chinese and Southern Han Chinese exhibit both Ancient Northern East Asian and Ancient Southern East Asian ancestries.[103] These genetic observations are in line with historical records of continuous and large migratory waves of northern China inhabitants escaping warfare and famine, to southern China. Aside from these large migratory waves, other smaller southward migrations occurred during almost all periods in the past two millennia.[104] A study by the Chinese Academy of Sciences into the gene frequency data of Han subpopulations and ethnic minorities in China showed that Han subpopulations in different regions are also genetically quite close to the local ethnic minorities, suggesting that in many cases, ethnic minorities ancestry had mixed with Han, while at the same time, the Han ancestry had also mixed with the local ethnic minorities.[105]

Han Chinese, similar to other East Asian populations, have inherited European ancestry, around 2.8% in Northern Han Chinese and around 1.7% in Southern Han Chinese.[69]

An extensive, genome-wide association study of the Han population in 2008, shows that geographic-genetic stratification from north to south has occurred and centrally placed populations act as the conduit for outlying ones.[106] Ultimately, with the exception in some ethnolinguistic branches of the Han Chinese, such as Pinghua, there is "coherent genetic structure" (homogeneity) in all Han Chinese.[107]

Paternal lineages

The major haplogroups of Han Chinese belong to subclades of Haplogroup O-M175. Y-chromosome O2-M122 is a common DNA marker in Han Chinese, as it appeared in China in prehistoric times, and is found in more than 50% of Chinese males, with frequencies tending to be high toward the east of the country, ranging from 29.7% to 52% in Han from southern and central China, to 55–68% in Han from the eastern and northeastern Chinese mainland and Taiwan.[108]

Other Y-DNA haplogroups that have been found with notable frequency in samples of Han Chinese include

O-M268(xM95, M176) (4.7–7%), and Q-M242 (2/168 = 1.2–4.2%).[109][108]

Maternal lineages

The mitochondrial-DNA haplogroups of the Han Chinese can be classified into the northern East Asian-dominating haplogroups, including A, C, D, G, M8, M9, and Z, and the southern East Asian-dominating haplogroups, including B, F, M7, N*, and R.[104]

These haplogroups account for 52.7% and 33.85% of those in the Northern Han, respectively. Haplogroup D is the modal mtDNA haplogroup among northern East Asians. Among these haplogroups, D, B, F, and A were predominant in the Northern Han, with frequencies of 25.77%, 11.54%, 11.54%, and 8.08%, respectively.

However, in the Southern Han, the northern and southern East Asian-dominating mtDNA haplogroups accounted for 35.62% and 51.91%, respectively. The frequencies of haplogroups D, B, F, and A reached 15.68%, 20.85%, 16.29%, and 5.63%, respectively.[101][110][111][112][113]

Tibetan peoples

Main article: Tibetan people

The ethnic roots of Tibetans can be traced back to an deep Eastern Asian lineage representing the indigenous population of the Tibetan plateau since c. 40,000 to 30,000 years ago, and arriving Neolithic farmers from the Yellow River within the last 10,000 years associated, and which can be associated with having introduced the Sino-Tibetan languages. Modern Tibetans derive up to 20% from Paleolithic Tibetans, with the remaining 80% being primarily derived from Yellow River farmers. The present-day Tibetan gene pool was formed at least 5,100 years BP.[114][115]

Paternal lineage

Tibetan males predominantly belong to the paternal lineage D-M174 followed by lower amounts of O-M175.[116]

Maternal lineage

Tibetan females belong mainly to the Northeast Asian maternal haplogroups M9a1a, M9a1b, D4g2, D4i and G2ac, showing continuity with ancient middle and upper Yellow River populations.[117]

Turkic peoples

See also: Turkic peoples

Linguistic and genetic evidence strongly suggests an early presence of Turkic peoples in eastern Mongolia.[118] The genetic evidence suggests that the Turkification of Central Asia was carried out by East Asian dominant minorities migrating out of Mongolia.[29]

Genetic data found that almost all modern Turkic-speaking peoples retained at least some shared ancestry associated with "Southern Siberian and Mongolian" (SSM) populations, supporting this region as the "Inner Asian Homeland (IAH) of the pioneer carriers of Turkic languages" which subsequently expanded into Central Asia.[119]

Population structure of Turkic-speaking populations in the context of their geographic neighbors across Eurasia. Turkic-speaking populations are shown in red. The upper barplot shows only Turkic-speaking populations.
Genetic, archeologic and linguistic evidence links the early Turkic peoples with Northeast Asian millet-agriculturalists, which later adopted a nomadic lifestyle and expanded from eastern Mongolia westwards.

An Ancient Northeast Asian origin of the early Turkic peoples has been corroborated in multiple recent studies. Early and medieval Turkic groups however exhibited a wide range of both (Northern) East Asian and West Eurasian genetic origins, in part through long-term contact with neighboring peoples such as Iranian, Mongolic, Tocharian, Uralic and Yeniseian peoples, and others.[120][121][122][123][124][125]

Paternal lineages

Common Y-DNA haplogroups in Turkic peoples are

Nogais, Kazakhs, and Uzbeks).[126][127]

Relationship to other Asia-Pacific and Native American populations

Central Asians

See also: History of Central Asia § Medieval
PCA of various populations in the context of Eurasia and the Americas

The genetic evidence suggests that the Turkification of Central Asia was carried out by East Asian dominant minorities migrating out of Mongolia.[128] According to a recent study, the Turkic Central Asian populations, such as Kyrgyz, Kazakhs, Uzbeks, and Turkmens share more of their gene pool with various East Asian and Siberian populations than with West Asian or European populations. The study further suggests that both migration and linguistic assimilation helped to spread the Turkic languages in Eurasia.[129]

North Asians and Native Americans

Genetic data suggests that

Ancient North Eurasians
or Ancient North Siberians.

Between 30,000 and 25,000 years ago, the ancestors of both Paleo-Siberians and Native Americans originated from admixture between

Ancient North Eurasians/Siberians and an Ancient East Asian lineage.[130][131] Ancestral Native Americans (or Ancient Beringians) later migrated towards the Beringian region, became isolated from other populations, and subsequently populated the Americas. Further geneflow from Northeast Asia resulted in the modern distribution of "Neo-Siberians" (associated with 'Altaic speakers') through the merger of Paleo-Siberians with Northeast Asians.[132][133][134]

Overall, while Northern Asians cluster closely to East Asians, they are shifted into a distinct position. "Analyses of all 122 populations confirm many known relationships and show that most populations from North Asia form a cluster distinct from all other groups. Refinement of analyses on smaller subsets of populations reinforces the distinctiveness of North Asia and shows that the North Asia cluster identifies a region that is ancestral to Native Americans."[135]

Native Americans

Multiple studies suggests that all Native Americans ultimately descended from a single founding population that initially diverged from" Ancestral Beringians" which shared a common origin with Paleo-Siberians from the merger of Ancient North Eurasians and a Basal-East Asian source population in Mainland Southeast Asia around 36,000 years ago, at the same time at which the proper

Paleosiberians, confirming that Ancestral Native Americans split from an ancient Siberian source population somewhere in northeastern Siberia. Genetic data on samples with alleged "Paleo-Indian" morphology turned out to be closely related to contemporary Native Americans, disproving a hypothetical earlier migration into the Americas. The scientists suggest that variation within Native American morphology is just that, the natural variation which have arisen during the formation of Ancestral Native Americans. Signals of a hypothetical "population Y", if not a false positive, are likely explained through a now extinct population from East Asia (e.g. Tianyuan man, which contributed low amounts of ancestry to the Ancestral Native American gene pool in Asia, and perhaps also towards other Asian and Oceanian populations.[137][136][138][139][140]

South Asians

Main article:
Genetic history of South Asia

The genetic makeup of modern South Asians can be described as a combination of West Eurasian ancestries with divergent

Himalayan foothills and Northeast India,[142][143] and is generally distributed throughout South Asia at lower frequency, with substantial presence in Mundari-speaking groups.[142][143]

According to a genetic research (2015) including linguistic analyses, suggests an East Asian origin for proto-Austroasiatic groups, which first migrated to Southeast Asia and later into India.[144] According to Ness, there are three broad theories on the origins of the Austroasiatic speakers, namely northeastern India, central or southern China, or southeast Asia.[145] Multiple researches indicate that the Austroasiatic populations in India are derived from (mostly male dominated) migrations from Southeast Asia during the Holocene.[146][144][147][148][149][150] According to Van Driem (2007), "...the mitochondrial picture indicates that the Munda maternal lineage derives from the earliest human settlers on the Subcontinent, whilst the predominant Y chromosome haplogroup argues for a Southeast Asian paternal homeland for Austroasiatic language communities in India."[147]: 7 

According to Chaubey et al. (2011), "Austroasiatic speakers in India today are derived from dispersal from Southeast Asia, followed by extensive sex-specific admixture with local Indian populations."[146] According to Zhang et al. (2015), Austroasiatic (male) migrations from southeast Asia into India took place after the lates Glacial maximum, circa 4,000 years ago.[144] According to Arunkumar et al. (2015), Y-chromosomal haplogroup O2a1-M95, which is typical for Austroasiatic speaking peoples, clearly decreases from Laos to east India, with "a serial decrease in expansion time from east to west," namely "5.7 ± 0.3 Kya in Laos, 5.2 ± 0.6 in Northeast India, and 4.3 ± 0.2 in East India." This suggests "a late Neolithic east to west spread of the lineage O2a1-M95 from Laos."[149][151] According to Riccio et al. (2011), the Munda people are likely descended from Austroasiatic migrants from southeast Asia.[148][152] According to Ness, the Khasi probably migrated into India in the first millennium BCE.[145]

According to Yelmen et al. 2019, the two main components of Indian genetic variation; the South Asian populations that "separated from East Asian and Andamanese populations" form one of the deepest splits among non-African groups compared to the West Eurasian component because of "40,000 years of independent evolution".[153]

Geneflow from Southeast Asians (particularly Austroasiatic groups) to South Asian peoples is associated with the introduction of rice-agriculture to South Asia. There is significant cultural, linguistic, and political Austroasiatic influence on early India, which can also be observed by the presence of Austroasiatic loanwords within Indo-Aryan languages.[154][155]

Southeast Asians

Estimated ancestry components among selected modern populations per Changmai et al. (2022). The yellow component represents East Asian-like ancestry.[156]

A 2020 genetic study about Southeast Asian populations, found that mostly all Southeast Asians are closely related to East Asians and have mostly "East Asian-related" ancestry.[18][157]

Ancient remains of hunter-gatherers in Maritime Southeast Asia, such as one Holocene hunter-gatherer from South Sulawesi, had ancestry from both, an Australasian lineage (represented by Papuans and Aboriginal Australasians) and an "Ancient Asian" lineage (represented by East Asians or Andamanese Onge). The hunter-gatherer individual had approximately c. 50% "Basal-East Asian" ancestry and c. 50% Australasian/Papuan ancestry, and was positioned in between modern East Asians and Papuans of Oceania. The authors concluded that East Asian-related ancestry expanded from Mainland Southeast Asia into Maritime Southeast Asia much earlier than previously suggested, as early as 25,000 BCE, long before the expansion of Austroasiatic and Austronesian groups.[158]

A 2022 genetic study confirmed the close link between East Asians and Southeast Asians, which the authors term "East/Southeast Asian" (ESEA) populations, and also found a low but consistent proportion of South Asian-associated "SAS ancestry" (best samplified by modern

Dhaka, Bangladesh) among specific Mainland Southeast Asian (MESA) ethnic groups (~2–16% as inferred by qpAdm), likely as a result of cultural diffision; mainly of South Asian merchants spreading Hinduism and Buddhism among the Indianized kingdoms of Southeast Asia. The authors however caution that Bengali samples harbor detechtable East Asian ancestry, which may affect the estimation of shared haplotypes. Overall, the geneflow event is estimated to have happened between 500 and 1000 YBP.[159]

Australasians

Main articles: Indigenous people of New Guinea, Melanesians, and Aboriginal Australians
PCA plot of genetic variationof worldwide populations. Australasians (green) cluster relative close to other East Eurasians, such as East/Southeast Asians.

Melanesians and Aboriginal Australians are deeply related to East Asians. Genetic studies have revealed that Australasians descended from the same Eastern Eurasian source population as East Asians and indigenous South Asians (AASI).[3]

Notes

References

  1. PMID 26530905
    .
  2. PMID 35445261
    .
  3. ^
    ISSN 2770-5005
    . ...In contrast, mainland East and Southeast Asians and other Pacific islanders (e.g., Austronesian speakers) are closely related to each other [9,15,16] and here denoted as belonging to an East and Southeast Asian (ESEA) lineage (Box 2). …the ESEA lineage differentiated into at least three distinct ancestries: Tianyuan ancestry which can be found 40,000–33,000 years ago in northern East Asia, ancestry found today across present-day populations of East Asia, Southeast Asia, and Siberia, but whose origins are unknown, and Hòabìnhian ancestry found 8,000–4,000 years ago in Southeast Asia, but whose origins in the Upper Paleolithic are unknown.
  4. ^ Genetics and material culture support repeated expansions into Paleolithic Eurasia from a population hub out of Africa, Vallini et al. 2022 (4 April 2022) Quote: "Taken together with a lower bound of the final settlement of Sahul at 37 ka (the date of the deepest population splits estimated by Malaspinas et al. 2016), it is reasonable to describe Papuans as either an almost even mixture between East Asians and a lineage basal to West and East Asians occurred sometimes between 45 and 38 ka, or as a sister lineage of East Asians with or without a minor basal OoA or xOoA contribution. We here chose to parsimoniously describe Papuans as a simple sister group of Tianyuan, cautioning that this may be just one out of six equifinal possibilities."
  5. ^ "Almost all living people outside of Africa trace back to a single migration more than 50,000 years ago". science.org. Retrieved 19 August 2022.
  6. ^
    S2CID 234247309
    . Via the southern route, ancestors of current Asian populations reached Southeast Asia and a part of Oceania around 70000–50000 years ago, probably through a coastal dispersal route (Bae et al., 2017). The oldest samples providing the genetic evidence of the northern migration route come from a high-coverage genome sequence of individuals excavated from the Yana RHS site in northeastern Siberia (Figure 2), which is about 31600 years old (Sikora et al., 2019). A wide range of artifacts, including bone crafts of wooly rhinoceros and mammoths, were excavated at the site (Pitulko et al., 2004). The analysis of genome sequences showed that the samples were deeply diverged from most present-day East Asians and more closely related to present-day Europeans, suggesting that the population reached the area through a route different from the southern route. A 24000-year-old individual excavated near Lake Baikal (Figure 2), also known as the Mal'ta boy, and 17000-year-old individuals from the Afontova Gora II site (Afontova Gora 2 and 3) showed similar genetic features to the Yana individuals (Raghavan et al., 2014; Fu et al., 2016; Sikora et al., 2019). Interestingly, genetic data suggested that Yana individuals received a large amount of gene flow from the East Asian lineage (Sikora et al., 2019; Yang et al., 2020).
  7. PMID 32843717
    . Population genomic studies on present-day humans7,8 have exclusively supported the southern route origin of East Asian populations.
  8. PMID 33753512
    .
  9. S2CID 4436271
    .
  10. PMID 37644833
    . A single major migration of modern humans into the continents of Asia and Sahul was strongly supported by earlier studies using mitochondrial DNA, the non-recombining portion of Y chromosomes, and autosomal SNP data [42–45]. Ancestral Ancient South Indians with no West Eurasian relatedness, East Asians, Onge (Andamanese hunter–gatherers) and Papuans all derive in a short evolutionary time from the eastward dispersal of an out-of-Africa population [46,47], although Europeans and East Asians are suggested to share more recent common ancestors than with Papuans [48]. The HUGO (Human Genome Organization) Pan-Asian SNP consortium [44] investigated haplotype diversity within present-day Asian populations and found a strong correlation with latitude, with diversity decreasing from south to north. The correlation continues to hold when only mainland Southeast Asian and East Asian populations are considered, and is perhaps attributable to a serial founder effect [50]. These observations are consistent with the view that soon after the single eastward migration of modern humans, East Asians diverged in southern East Asia and dispersed northward across the continent.
  11. PMID 22908291
    . Inferences from nuclear (51), Y chromosome (52), and mitochondrial genome (53) data support an early migration of modern humans out of Africa and into Southeast Asia using a southern route by at least 60 ka. Patterns of genetic variation in recent human populations (11, 54, 55) recognize Southeast Asia as an important source for the peopling of East Asia and Australasia via a rapid, early settlement.
  12. ISBN 978-981-10-4169-3
    . "In the study of Zhong et al. haplogroups O-M175, C-M130, D-M174 and N-M231 still suggests the substantial contribution of the southern route. However, the Central Asia and West Eurasia related haplogroups, such as haplogroups R-M207 and Q-M242, occur primarily in northwestern East Asia and their frequencies gradually decrease from west to east. In addition, the Y-STR diversities of haplogroups R-M207 and Q-M242 also indicate the existence of northern route migration about 18,000 years ago from Central Asia to North Asia, and recent population admixture along the Silk Road since about 3000 years ago (Piazza 1998)."
  13. S2CID 250502011
    . "In addition to the earliest southern settlement of AMHs in East Asia, ancient migration (40–18 kya) into East Asia via the “Northern Route” from West Eurasia was previously proposed. The “Northern Route” hypothesis would also explain where the subtle shared ancient north Eurasian (ANE) ancestry came from that is then also shared with Native Americans."
  14. PMID 34410389
    .
  15. PMID 23731529
    .
  16. PMID 27965293
    .
  17. S2CID 235226413
    .
  18. ^
    PMID 33781248
    .
  19. ^ "Direct genetic evidence of founding population reveals story of first Native Americans". University of Cambridge. 3 January 2018. Retrieved 9 June 2021.
  20. PMID 32344428
    .
  21. bioRxiv 10.1101/2020.12.07.414037
    .
  22. S2CID 218926428
    .
  23. PMID 15339343
    .
  24. ^
    S2CID 220671047
    .
  25. ^ "Xiongnu People". britannica.com. Encyclopædia Britannica. Archived from the original on 11 March 2020. Retrieved 25 July 2015.
  26. ^ di Cosmo 2004: 186
  27. ^
    S2CID 218935871. Text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
    . "Such a distribution of Xiongnu words may be an indication that both Turkic and Eastern Iranian-speaking groups were present among the Xiongnu in the earlier period of their history. Etymological analysis shows that some crucial components in the Xiongnu political, economic and cultural package, including dairy pastoralism and elements of state organization, may have been imported by the Eastern Iranians. Arguably, these Iranian-speaking groups were assimilated over time by the predominant Turkic-speaking part of the Xiongnu population. ... The genetic profile of published Xiongnu individuals speaks against the Yeniseian hypothesis, assuming that modern Yeniseian speakers (i.e. Kets) are representative of the ancestry components in the historical Yeniseian speaking groups in southern Siberia. In contrast to the Iron Age populations listed in Table 2, Kets do not have the Iranian-related ancestry component but harbour a strong genetic affinity with Samoyedic-speaking neighbours, such as Selkups (Jeong et al., 2018, 2019)."
  28. ^ Savelyev & Jeong 2020:"Specifically, individuals from Iron Age steppe and Xiongnu have an ancestry related to present-day and ancient Iranian/Caucasus/Turan populations in addition to the ancestry components derived from the Late Bronze Age populations. We estimate that they derive between 5 and 25% of their ancestry from this new source, with 18% for Xiongnu (Table 2). We speculate that the introduction of this new western Eurasian ancestry may be linked to the Iranian elements in the Xiongnu linguistic material, while the Turkic-related component may be brought by their eastern Eurasian genetic substratum." Table 2: Sintashta_MLBA, 0.239; Khovsgol LBA, 0.582; Gonur1 BA 0.178
  29. ^
    S2CID 13670282
    .
  30. PMID 37058560
    .
  31. ^ Rogers & Kaestle 2022.
  32. ^ Li H (2012). Y-Chromosome Genetic Diversity of the Ancient North Chinese populations (Report). China: Jilin University.
  33. PMID 37058560
    .
  34. ^ Lee 2023: Haplogroup information for DA39 is located in Supplementary Materials document Data S1 (abk1900_Data_S1.xlsx), row 58. "Before this study, only one other individual from an elite square tomb had been analyzed in a genome-wide manner: DA39 from Tomb 1 at the imperial elite site of Gol Mod 2 in central-north Mongolia (13). This adult male, buried in one of the largest square tomb complexes excavated to date, surrounded by at least 27 satellite burials, and containing rare exotic items such as Roman glass bowls, was likely a chanyu, or ruler of the empire (73, 74)."
  35. S2CID 246508594
    . " The first pattern is that the slab burial mtDNA frequencies are extremely similar to those of the aggregated Xiongnu populations and relatively similar to those of the various Bronze Age Mongolian populations, strongly supporting a population continuity hypothesis for the region over these time periods (Honeychurch, 2013)"
  36. PMID 12858290
    .
  37. S2CID 165623743
    . "Analysis of the mitochondrial DNA, which is maternally inherited, shows that the Xiongnu remains from this Egyin Gol necropolis consist mainly of Asian lineages (89%). West Eurasian lineages makeup the rest (11%) (Keyser-Tracqui et al. (2003: 258). However, according to a more recent study of ancient human remains from central Mongolia, the Xiongnu population in central Mongolia possessed a higher frequency of western mitochondrial DNA haplotypes (37.5%) than the Xiongnu from the Egyin Gol necropolis (Rogers 2016: 78)."
  38. S2CID 265381985
    .
  39. ^ Li et al. 2018, p. 1.
  40. S2CID 19492267
    .
  41. PMID 17632796
    .
  42. S2CID 18809679
    .
  43. doi:10.1537/ase.201215
    .
  44. ^
    PMID 34533991
    .
  45. S2CID 206667111
    .
  46. doi:10.47248/hpgg2202010001
    .
  47. ISSN 1000-3193
    .
  48. ^
    PMID 32843717
    .
  49. PMID 34659368.  This article incorporates text available under the CC BY 4.0
    license.
  50. ^ a b c Yali Xue, Tatiana Zerjal, Weidong Bao, Suling Zhu, Qunfang Shu, Jiujin Xu, Ruofu Du, Songbin Fu, Pu Li, Matthew E. Hurles, Huanming Yang, and Chris Tyler-Smith, "Male Demography in East Asia: A North–South Contrast in Human Population Expansion Times." Genetics 172: 2431–2439 (April 2006). DOI: 10.1534/genetics.105.054270
  51. ^ a b c Wang Chi-zao, Shi Mei-sen, and Li Hui, "The Origin of Daur from the Perspective of Molecular Anthropology." Journal of North Minzu University 2018, No．5, Gen．No．143.
  52. S2CID 7685248
    .
  53. S2CID 7505563
    .
  54. ^ a b "Did you know DNA was used to uncover the origin of the House of Aisin Gioro?". Did You Know DNA... 14 November 2016. Retrieved 5 November 2020.
  55. PMID 16380921
    .
  56. ^ "Asian Ancestry based on Studies of Y-DNA Variation: Part 3. Recent demographics and ancestry of the male East Asians – Empires and Dynasties". Genebase Tutorials. Archived from the original on 25 November 2013.
  57. PMID 34659368
    .
  58. ^ Michael F. Hammer, Tatiana M. Karafet, Hwayong Park, Keiichi Omoto, Shinji Harihara, Mark Stoneking, and Satoshi Horai, "Dual origins of the Japanese: common ground for hunter-gatherer and farmer Y chromosomes." J Hum Genet (2006) 51:47–58. DOI 10.1007/s10038-005-0322-0
  59. ^ "公益財団法人 アイヌ民族文化財団". www.ff-ainu.or.jp (in Japanese). Retrieved 8 December 2023.
  60. ISBN 9784585053941
    .
  61. ^ a b Suzuki, Yuka (6 December 2012). "Ryukyuan, Ainu People Genetically Similar". Asian Scientist. Archived from the original on 16 June 2016. Retrieved 11 June 2016.
  62. PMID 34533991
    .
  63. ^ "'Jomon woman' helps solve Japan's genetic mystery | NHK WORLD-JAPAN News". NHK WORLD. Retrieved 19 November 2020.
  64. ^ 弥生人ＤＮＡで迫る日本人の起源」 [The origin of Japanese people approaching with Yayoi DNA]. ja:サイエンスZERO (Television production) (in Japanese). NHK. 23 December 2018.
  65. PMID 37588377
    .
  66. PMID 33618348
    .
  67. PMID 33618348
    .
  68. ^ Nikkei Science (23 June 2021). "渡来人、四国に多かった？ ゲノムが明かす日本人ルーツ" [Were there many migrants in Shikoku? Japanese roots revealed by genome analysis]. nikkei.com (in Japanese). Retrieved 1 May 2022.
  69. ^
    PMID 25833680
    .
  70. S2CID 12893406
    .
  71. ^ The JPT sample is considered "as generally representative of the majority population in Japan". See Matsuda I. "Japanese in Tokyo, Japan – Population Description". Camden, NJ: Coriell Institute for Medical Research.
  72. PMID 27111036
    .
  73. PMID 21998705
    .
  74. ^
    PMID 19148289
    .
  75. ^ Reference Populations – Geno 2.0 Next Generation. (2017). The Genographic Project. Retrieved 15 May 2017, from link.
  76. ^
    PMID 21463511. Text was copied from this source, which is available under a Creative Commons Attribution 2.0 (CC BY 2.0)
    license.
  77. ^
    S2CID 251410856
    .
  78. ^ Wang & Wang 2022: "The genetic legacy of Jomon was not restricted to Japan but was also found in Neolithic Korea."
  79. ^ Wang & Wang 2022: "Taking no account of this Bronze Age sample, the possible scenario was that the indigenous Jomon-related ancestry was largely replaced through admixture with the incoming northern China populations and disappeared in the later centuries, creating relatively homogeneous present-day Koreans."
  80. ^ Wang & Wang 2022: "Present-day Koreans are a highly homogeneous population without apparent genetic substructure on a whole-genome scale and with close genetic relationships with Japanese and northern Han Chinese. [...] The genetic variation of the Y chromosome and mitochondrial DNA show that Koreans contain lineages typical of both Southeast and Northeast Asian populations, suggesting that Korea was populated in multiple waves"
  81. PMC 9827920
    . "In both ancient and present-day Koreans, we do not detect a statistically significant contribution from the Jomon hunter-gatherer gene pool of the Japanese archipelago (Table S7A), although previous studies report occasional presence of the Jomon ancestry contribution from Neolithic to the early Medieval period (Gelabert et al., 2022; Robbeets et al., 2021). When we replace the genetic northern proxy from WLR_BA to Middle Neolithic individuals from the Miaogizou site in Inner Mongolia (“Miaozigou_MN”), we detect a small but significant amount of Jomon contribution in the Gunsan individuals and present-day Ulsan Koreans (3.1%–4.4%; Table S7B). We believe that WLR_BA provides a more suitable model for ancient and present-day Koreans given its geographical and temporal proximity to them. The remaining well-fitting source pairs provide qualitatively similar results (Table S8)."
  82. S2CID 27125545
    .
  83. ^ a b c d e So Yeun Kwon, Hwan Young Lee, Eun Young Lee, Woo Ick Yang, and Kyoung-Jin Shin, "Confirmation of Y haplogroup tree topologies with newly suggested Y-SNPs for the C2, O2b and O3a subhaplogroups." Forensic Science International: Genetics 19 (2015) 42–46. http://dx.doi.org/10.1016/j.fsigen.2015.06.003
  84. doi:10.7580/KoreanJLegMed.2012.36.1.34
    .
  85. ISSN 1875-1768
    .
  86. S2CID 27125545
    .
  87. PMID 19148289
    .
  88. PMID 32766443
    .
  89. S2CID 256067815
    .
  90. ^ "Population dynamics and the rise of empires in Inner Asia: Genome-wide analysis spanning 6,000 years in the eastern Eurasian Steppe gives insights to the formation of Mongolia's empires". ScienceDaily. Retrieved 4 June 2023.
  91. PMID 34956310
    .
  92. ^ "DNA Match Solves Ancient Mystery". china.org.cn. Retrieved 25 December 2018.
  93. S2CID 13217444
    .
  94. ^ Guang‐Lin He, Meng‐Ge Wang, Xing Zou, Hui‐Yuan Yeh, Chang‐Hui Liu, Chao Liu, Gang Chen, and Chuan‐Chao Wang. "Extensive ethnolinguistic diversity at the crossroads of North China and South Siberia reflects multiple sources of genetic diversity." J Syst Evol, 2023, 61(1): 230-250. https://doi.org/10.1111/jse.12827
  95. ^ "Mongol Genetics – DNA of Mongolia's Khalkha Mongolians and others". khazaria.com. Retrieved 25 December 2018.
  96. PMID 8846908
    .
  97. S2CID 218649510
    .
  98. JSTOR 48726991
    .
  99. PMID 29636655
    .
  100. PMID 32355288
    .
  101. ^
    PMID 25938511
    .
  102. PMID 29202706
    .
  103. S2CID 218649510
    .
  104. ^
    S2CID 4301581
    .
  105. S2CID 1924085
    .
  106. PMID 19944401
    .
  107. PMID 18270655
    .
  108. ^
    PMID 36973821
    .
  109. PMID 19088127
    .
  110. PMID 11836649
    .
  111. PMID 12270900
    .
  112. PMID 12598688
    .
  113. PMID 20713468
    .
  114. S2CID 247317520
    .
  115. PMID 36930720
    .
  116. PMID 26538459
    . "Comparing Sherpas, Tibetans, and Han Chinese showed that the D-M174 is the predominant haplogroup in Sherpas (43.38%) and prevalent in Tibetans (52.84%)5, but rare among both Han Chinese (1.4–6.51%)6,7 and other Asian populations (0.02–0.07%)8, aside from Japanese (34.7%) who possesses a distinct D-M174 lineage highly diverged from those in Tibetans and other Asian populations9,10."
  117. S2CID 257588399
    .
  118. ISBN 978-1-134-82869-2
    .
  119. PMID 25898006
    .
  120. S2CID 213436897
    .
  121. S2CID 213952845
    .
  122. S2CID 219470000
    .
  123. S2CID 166026934
    .
  124. PMID 36006373
    .
  125. PMID 35031610
    .
  126. PMID 12145751
    .
  127. ISBN 978-0-19-970547-4
    .
  128. S2CID 13670282
    .
  129. PMID 25898006
    .
  130. ISBN 978-1-108-49822-7
    .
  131. ^ Meltzer 2021, p. 170: "The ancestors of Native Americans derive from descendants of Ancient North Siberians and Ancient North Eurasians on the one hand, (exemplified by the genomes of the Yana and Mal'ta individuals, respectively), and a population of Ancient East Asians (signified by the Tianyuan individual). From those groups emerged the deepest ancestors of Native Americans, who became separated far in far northeastern Asia from their Siberian and Asian ancestors around the time of the LGM. Part of the population stayed in Siberia (the Ancient Paleo-Siberians, seen in the Kolyma genome), while others moved east across Beringia."
  132. ^ Lazaridis et al. 2018.
  133. ^ Moreno-Mayar et al. 2018.
  134. PMID 29033327
    .
  135. PMID 35508562
    .
  136. ^
    S2CID 4454580
    .
  137. PMID 31467216
    .
  138. PMID 30415837
    .
  139. S2CID 235460793
    .
  140. ^ Sarkar AA (18 June 2021). "Ancient Human Genomes Reveal Peopling of the Americas". GEN – Genetic Engineering and Biotechnology News. Retrieved 15 September 2021. The team discovered that the Spirit Cave remains came from a Native American while dismissing a longstanding theory that a group called Paleoamericans existed in North America before Native Americans.
  141. PMID 30952160
    .
  142. ^ a b Chaubey G (January 2015). "East Asian ancestry in India" (PDF). Indian Journal of Physical Anthropology and Human Genetics. 34 (2): 193–199. Here the analysis of genome wide data on Indian and East/Southeast Asian demonstrated their restricted distinctive ancestry in India mainly running along the foothills of Himalaya and northeastern part.
  143. ^
    PMID 20978040
    .
  144. ^
    PMID 26482917
    .
  145. ^
    ISBN 978-1-118-97058-4
    .
  146. ^
    PMID 20978040
    .
  147. ^ a b Van Driem G (2007). "Austroasiatic phylogeny and the Austroasiatic homeland in light of recent population genetic studies" (PDF). Mon-Khmer Studies. 37: 1–14.[dead link]
  148. ^
    S2CID 39428816
    .
  149. ^
    S2CID 83103649
    .
  150. ^ "ASI-AAA"
  151. ^ Valar M (21 April 2015). "DNA Reveals Unknown Ancient Migration Into India". National Geographic. Archived from the original on 11 June 2015.
  152. ^ Gutman A, Avanzati B. "Austroasiatic Languages". The Language Gulper.
  153. PMID 30952160
    .
  154. ISBN 978-81-206-0772-9
    . It has been further proved that not only linguistic but also certain cultural and political facts of ancient India, can be explained by Austroasiatic elements.
  155. ^ "How rice farming may have spread across the ancient world". science.org. Retrieved 26 October 2021.
  156. PMID 36581666
    .
  157. PMID 32344428
    .
  158. PMID 34433944
    . The qpGraph analysis confirmed this branching pattern, with the Leang Panninge individual branching off from the Near Oceanian clade after the Denisovan gene flow, although with the most supported topology indicating around 50% of a basal East Asian component contributing to the Leang Panninge genome (Fig. 3c, Supplementary Figs. 7–11).
  159. PMID 35176016
    .

Works cited

Further reading

Retrieved from "https://en.wikipedia.org/w/index.php?title=Genetic_history_of_East_Asians&oldid=1220984594"