Who are the Ainu people?

Ainu man circa 1880

The Ainu are an indigenous ethnic group of people who live in Hokkaido in Japan today as well as in Russia (the Kuril Islands and Sakhalin). In the 19th Century, Japanese people called the northern island of Hokkaido “Ezochi” which means “Land of the Ainu”. The term Ainu generally referred to the fair-skinned, long-haired hunter-gatherer-fishering people with animistic beliefs who had lived there for hundreds of years.

From the 15th century, waves of Japanese settlers began crowding out Ainu communities on Honshu island and pushing them northwards. The settlers also brought infectious diseases that caused Ainu populations to fall. Ainu land was redistributed to Japanese farmers.

Map showing location of Ainu populations

1899年,日本政府通过了开云体彩官网一项法案which labelled the Ainu “former Aborigines”, ostensibly declaring that the Ainu had been integrated into the Japanese population – the act, together with the various assimilation policies had the drastic effect of eroding Ainu identity and traditions. The Meiji government’s 1899 assimilation policies resulted in the ban of the Ainu language and Ainu children being given Japanese names and put into Japanese schools. As a result of these policies, many Ainu people suffered discrimination and became ashamed of their language and culture. The act continued for a hundred years.

The 1899 act was finally officially reversed on June 6th, 2008, when the Japanese government passed a resolution adopt a resolution that, for the first time, formally recognised the Ainu as “an indigenous people who have their own language, religion and culture”.

今天只剩下少量的阿伊努人,他们constitute one of Japan’s most marginalised groups. The Ainu are thought to number around 25,000 (official sources) while unofficially, they are believed to number around 200,000 or more since many Ainu still do not disclose their roots out of fear of discrimination.

Origins: Where did the Ainu come from?

Historically, they spoke the Ainu language and have traditionally been considered the descendents of the Jomon or post-Jomon people of Japan. In their Yukar Upopo (Ainu Legends) is told, “The Ainu lived in this place a hundred thousand years before the Children of the Sun came”.

The enduring puzzle of the origin of the unique D2 Y-DNA of the Ainu

Groundbreaking genetic mapping studies by Cavalli-Sforza have shown a sharp gradient in gene frequencies centered in the area around the Sea of Japan, suggesting that the area was a center of expansion for the ancestral Jomon-Ainu populations (thought to have occurred during the Jomon period although the studies cannot fix clear dates). This expansion of populations is thought to be the third most important genetic movement in Eurasia (after the “Great expansion” from the African continent, to Arabia and adjacent parts of the Middle East, as well as to the northern regions of Eurasia, (particularly Siberia from regions to the south).

According to genetic tests, the Ainu people belong mainly to Y-DNA haplogroupD2(a haplogroup that is found uniquely in and frequently throughout Japan including Okinawa with itsclosest relations being Tibetans and Andaman Islanders in the Indian Ocean). On the paternal side, the vast majority (87.5%) of the Ainu were, according to a 2004 studyto be of Asian-specific YAP+ lineages (Y-haplogroups D-M55* and D-M125), that were only distributed in the Japanese Archipelago. The Ainu exhibited no other Y-haplogroups (i.e. none of the common East Asian C-M8, O-M175*, and O-M122* haplogroups) and shared no other Y-DNA in common in mainland Japanese and Okinawans.

According toa study led by Hammer, one of the most useful and widely studied Y-linked polymorphisms is known as the “Y Alu polymorphic” (YAP) element (Hammer 1994). This polymorphism has resulted from the single and stable insertion of a member of the repetitive Alu family at a specific site (locus DYS287) on the long arm of the human Y chromosome during the past 29,000-334,000 years.

The frequency of Y chromosomes carrying the YAP element (YAP+) varies greatly among human populations from different geographic locations: Global surveys have shown that sub-Saharan African populations have the highest overall frequency of YAP chromosomes, followed by populations from northern Africa, Asia, Europe, the New World, and Oceania. However, an intriguingfinding by Hammer (1997)that the ancestral YAP haplotype is the Asian haplotype 3 from which other haplotypes 4 and 5 evolved and derived, suggesting the possibility that YAP haplotype 3 originated in Asia and migrated to Africa. This hypothesis is supported by the finding of high frequencies of haplotype 3 in some Asian populations (i.e., -50% in Tibet) and by the observation of higher levels of diversity (based on the number and frequency of alleles at the DYS1 9 microsatellite locus) associated with Asian versus African haplotype 3 chromosomes.

Chandrasekhar et al. 2007, also argued for the Asian origin of the YAP+ on the basis of evidence from the presence of the YAP insertion in Northeast Indian tribes and Andaman Islanders with haplogroup D that suggests that some of the M168 chromosomes gave rise to the YAP insertion and M174 mutation in South Asia. Others such asUnderhillandBravistand by an African origin for YAP+. The prevalence of the YAP+ allele in central Asian populations was alternatively suggested by some (Altheide and Hammer 1997; Jin and Su 2000; Karafet et al. 2001) to point toa genetic contribution to the east Asian populations from the northwest, probably from central Asia.

In Japan, the frequency of the YAP element ranges from 33% in Shizuoka to 56% in Okinawa, with an intermediate frequency of 39% in Aomori. The frequency is significantly higher in Okinawa than in Shizuoka (Fisher’s exact test, P = .0284), but the Okinawa frequency is not significantly different from the Aomori frequency (P= .2196). However, the frequency in Okinawa is significantly higher than in the combination of the two Honshu prefectures (P = .0256). 87.5% of the Ainu were, according to a 2004 study to be of Asian-specific YAP+ lineages. By contrast, YAP was absent from Korean male samples. This result is consistent with previous surveys that showed the YAP element to be polymorphic in Japan but absent in other Asian and Oceanic populations. In terms of antiquity as well as the relationship of the different YAP+ lineages, another interesting conclusion was made bythe Hammer analysis: “All pairwise F., values were calculated on the basis of YAP allele frequencies in Japan and Taiwan, as well as in 13 other populations (Hammer 1994; Spurdle et al. 1994b). The neighbor-joining method was used to generate a clustering diagram (fig. 2). All Asian and Oceanic populations, except the Japanese, form a single group that is closely allied with the European populations. The greatest genetic distance is the one that separates these Eurasian populations from the Japanese and African populations. The Okinawan and Honshu populations are separated; the former population clusters in the middle of the African groups, and the latter population clusters between the African and Eurasian samples.”

Map showing the Yap+ allele occurrence and distribution (in grey), among the other lineages

In other tests of two out of a sample of sixteen – i.e. 12.5% of Ainu men were found to belong to Haplogroup C3, which is the most common Y-chromosome haplogroup among the indigenous populations of the Russian Far East and Mongolia. A separate test (a sample of four Ainu men) found that one in four Ainu men belonged to haplogroup C3.

The presence of theC3 haplogroup is believed to reflect the genetic influence of the nomadic Nivkhs people of northern Sakhalin Island, with whom the Ainu have long-standing cultural interactions.

Relationship to other populations

While Y-chromosome haplogroup markers D2 and mtDNA D4 and M7a (and M7a1) indicate that Ainu are related to other Japanese populations in the rest of Japan, the various mtDNA studies may indicate that Ainu men took wives from a variety of locations throughout Central Asia, Siberia and the Russian Far East.

East Asian genes

M7 has been detected so far in China, Vietnam, the West Siberian Mansi, Mongols and island Southeast Asia, apart from from the Korean peninsula and Japanese islands where the subclades expanded. Haplogroup M7a has been found in Southeast Asia-Taiwan, but mainly among Japanese and Ryukyuans.Haplogroup M7is seen ascharacteristically distributed in East Asiawhile M7a is regarded as its daughter group specific for (pre-Jomon to Jomon) Japanese populations (while M7b2 is specific for Korean populations). The 2002 study puts the estimated settlement times for M7a, M7b and M7c at between 6,000 and 18,000 years … although it suggests that the M7a and M7b pioneer settlers may have entered even earlier ~ 30,000 years at a time when the Yellow Sea had fallen dry and was more like a large lake , but that the populations became bottle-necked toward the Last Glacial Maximum (LGM). Beyond the ancestral M7, it has beennotedthat M7 nested subclades are not shuffled between the Koreans and Japanese, so the M7a, M7b and M7c starlike clades are thought to represent the post LGM resettlement process, with M7a in the area of the southern Japanese Sea. This settlement event would have been contemporary with the spread of microblades, e.g. of the Suyanggae-type and before the onset of the Jomon culture.

It has also been noted that the Ainu of today are not pure descendants of the Jomon, but rather from the Jomon-Yayoi mixture of the Satsumon/Emishi people. The Emishi were Jomon descendants with Yayoi assimilated cultural traits. It is believed from place names in Tohoku that the Emishi spoke the Ainu language as well. As the Yayoi people pushed north, it is thought that the Emishi people advanced on Hokkaido, infusing the Jomon culture in Hokkaido with an agrarian society and with metal-using traits from the Yayoi culture. CitingGary Crawford, Paula Nielsen writes in “Origins of the Ainu People of Northern Japan” that “the Satsumon culture recently discovered in Hokkaido was descended from the Tohoku Emishi of northeastern Honshu who migrated to Hokkaido, bringing a fused culture of the Middle Yayoi, along with the ancient physical traits of the Jomon”.

And although the Ainu of Japan have traditionally considered descendants of the Jomon or post-Jomon Satsumon people (indicated by the D or D2 gene marker), they have been found to carry the Y-chromosome DNA haplogroup C3 showing a paternal lineage from North Asia including Sakhalin.

mtDNA haplogroups Y – An origin in the neighbouring Amur tribes of the contiguous zones?

Genetic testing showed a varied picture of the maternal lines, with the presence ofmitochondrial DNA – haplogroup Y (21.6%), haplogroup M7a (and M7a1) (15.7%), haplogroup D – especially D4, and haplogroup G.

MtDNA haplogroup Yis thought to have been likely the genetic influence from the Nivkhs, (although it is also present in the Tungusic peoples, Koreans, Mongols (including Kalmyks and Buryats), Tajiks, Chinese and other Central Asians, South Siberian Turkic peoples (e.g. Tuvans, Todjins, Soyots). The presence ofY1 lineages (Y is restricted to Northeast Asians and Ainu) among the Ainu also points to the migration route, from Siberian populations to the northernmost populations of the Japanese islands (fitting well with the archaeological record) but poses separate events from the settlement by M7a and M7b peoples.

A2020 studyof the maritime tribes of the Northeast Pacific coast,

“identified the novel haplogroup N9b1 in Primorye, which impliesa link between a component of the Udegey ancestry and the Hokkaido Jomon. …

The Udegey group is found to consist of two mtDNA haplogroups: N9b and M7a2 (see Table 1 and Additional file 6: Figures S5 and Additional file 7: Figure S6). Aside from the Udegey originating in the villages of Gvasygi and Agzu in the Sikhote-Alin/Primorye region, we sampled the Udegey individuals who married into Ulchi and Nivkhi families dispersed along the reaches of the Lower Amur [22]. Haplogroup N9b is represented mainly by lineages of four major subhaplogroups: N9b1, N9b2, N9b3, and N9b4 [34,35,36].We identified a novel N9b1 mitogenome (MH807371) in one individual from Primorye (Agzu), hence expanding the established geographical scope of the N9b1 haplogroup and disclosing a link between a component of the Udegey ancestry and the Hokkaido Jomonfrom Japan [36,37,38]. The second prevalent haplogroup is M7a2a3a, which was detected in 8/46 (17.4%) of the Udegey samples (Table 1, Additional file 7: Figure S6). The Udegey, as well as the Hokkaido Jomon, lack subhaplogroup M7a1, which is the predominant subhaplogroup in modern Japanese and Korean populations “

The study also proposed:

mtDNA haplogroup Y (a descendant of haplogroup N9) is proposed to indicate matrilineal genetic continuity between late Pleistocene hunter-gatherer groups and present-day populations in the Far East[22, 25,26,27,28,29]. Within Siberia, the majority of contemporaryY1 carriers cluster into Y1a marked by the coding change m.7933A > G (aged ~ 10.6 kya), whereas Y1b and Y1c are confined to continental China, Japan, and Korea(Additional file 2: Figure S1). Accordingly, two offshoots arose from the Y1a founding haplotype for this haplogroup. On one side, the newly refinedY1a1 haplotype defined by m.12732 T > C is well represented in Tungusic-speaking groups (e.g., Evenki, Udegey),while the other side harbors a back mutation at np m.16189 T > C relative to the Reconstructed Sapiens Reference Sequence (RSRS). Sequence diversification within theY1a-m.16189 T > C! haplogroup is characteristic (at most) of the Nivkhi from Sakhalin. The updated network analysis includes 10 Y1a2 sequencesdefined by m.12397A > G, of which7 are new from the coastal Koryak.

… we extended the survey of major mitochondrial lineages dispersed across the Russian Far East. Several components may be delineated in this regard. The first component traces back to East Eurasian hunter-gatherers and represents lineages belonging to subdivisions of haplogroups N9b and M7a2. The second is well represented byY1a and G1b and points to the Lower Amur as the ancestral homelandfor this and other haplogroups. The third comprisesD4e5, which establishes an association between the Oroki and interior eastern Eurasian populations. Last,rare D4m2a mtDNA exhibited by modern Siberians may have roots in Primorye,at the eastern edge of the continent, rather than a South-Central Siberian source.

Influx of Siberian genes? G1b a marker of the Okhotsk tribes, and the Tokarev ancestor

HaplogroupG has been found most frequently among indigenous populations of easternmost Siberia, but it is also common in the Altai-Sayan region of southern Siberia, and Central Asia. According to a2002年的研究, D and G stem from a common source or node, butG1 is restricted to Northeast Siberiawhile G2a is highest among Central Asians (8.8%) and occurs at above 3% in Tibetans and Ainu.

Based on non-metric cranial traits studies, the Ainu people occupy an intermediate position between Jomon and Northeast Asians (the Okhotsk) on the one hand, and between and to a lesser extent the recent Hokkaido Ainu.Both genetic and cranial measurement studies support the recent consensus view that the Ainu originated in the merger of the Satsumon and Okhotsk cultures around 1200 CE.

The Ainu are also considered to occupy an intermediate position between Jomon and Northeast Asians on the one hand, and between Jomon and the Native Americans on the other. Thus studies on the variations among the three Ainu series from Sakhalin Island, the northeast Hokkaido coast, and central/south Hokkaido shown in the study, conclude that there has beenadmixture between the ancestors of recent Ainu and northern groups such as the Okhotsk peoplein the post-Jomon periods.

Sato’s 2009 studyof the Okhotsk mtDNA concluded that there was gene flow from the Sakhalin Okhotsk people into the Ainu. A2019 studyfound that theancestors of the Tokarev culture of the Kolymar basin, belonged to the G1b haplogroupand that the Tokarev culture (at the Spafaryev site) was probably associated with the migration of the migration of ancient Amur tribes to the north.

A 2007 study investigated the phylogenetic status of the Okhotsk people that were distributed in northern and eastern Hokkaido as well as southern Sakhalin during the fifth to the thirteenth centuries, DNA was carefully extracted from human bone and tooth remains excavated from archaeological sites, see 2007,Origins and genetic features of the Okhotsk people, revealed by ancient mitochondrial DNA analysis, Journal of Human Genetics 52(7):618-27 DOI – 10.1007/s10038-007-0164-z:

16 mtDNA haplotypes were identified from 37 individuals of the Okhotsk people. Of the 16 haplotypes found, 6 were unique to the Okhotsk people, whereas the other 10 were shared by northeastern Asian people that are currently distributed around Sakhalin and downstream of the Amur River. The phylogenetic relationships inferred frommtDNA sequences showed that the Okhotsk people were more closely related to the Nivkhi and Ulchi people among populations of northeastern Asia. In addition, theOkhotsk people had a relatively closer genetic affinity with the Ainu people of Hokkaido, and were likely intermediates of gene flow from the from the northeastern Asian people to the Ainu people.These findings support the hypothesis that the Okhotsk culture joined the Satsumon culture (direct descendants of the Jomon people) resulting in the Ainu culture,as suggested by previous archaeological and anthropological studies.

DNA updates – aDNA of Ainu of the Edo era:

A study of Edo era Ainu DNA throws light on regional differences in Ainu origins:

A2018 studyof mitochondrial DNA haplogroups of 94 Ainu individuals from the Edo era concluded that the Ainu were formed from the Hokkaido Jomon people, but subsequently underwent considerable admixture with adjacent populations; the study:

a. confirmed thatthe genetic influence of the Okhotsk culture people on the Ainu is significant. The proportion of Okhotsk‐type haplogroups in the Edo Ainu was 35.1%,which is as high as that of the Jomon‐type haplogroups (30.9%). This suggests that the Okhotsk culture people were one of the main genetic contributors to the formation of the Ainu.

b. indicated that while the Ainu still retain the matrilineage of the Hokkaido Jomon people, the Siberian influence on this population is far greater than previously recognized. Siberian‐type haplogroups are observed in the Edo Ainu. Although their frequency is low (7.3%), as described earlier, the existence of these haplogroups may hint at the continuity of the genetic relationship between the Ainu and native Siberians even after the Okhotsk culture disappeared from Hokkaido. However, the number of Okhotsk people who were genetically analyzed is still small (n = 37; Sato et al., 2009), so it is possible that these haplogroups will be identified in the Okhotsk people in further study.

c. ***It also found thatthe influence of mainland Japanese is evident PRIOR to the Meiji period, especially in the southwestern part of Hokkaidothat is adjacent to Honshu, the main island of Japan. Intriguingly,a genetic contribution of the mainland Japanese to the Edo Ainu is evident (28.1%), which is almost as considerableas those of the Jomon and the Okhotsk culture people. Conventionally, the genetic influence of the mainland Japanese on the Ainu is considered to have been limited until the Meiji government started sending settlers to Hokkaido as a national policy in 1869. However, our findings cast doubt on this accepted notion.

d. Regional differences of the Ainu were observed as follows (excerpt from study below):

By classifying the mtDNA haplogroups into four types as described earlier, regional differences of the Ainu people were highlighted. Judging from the data shown in Table 2, the high frequencies of Jomon‐type haplogroups in northeastern/central Hokkaido (44.2%) and the high frequencies of mainland Japanese‐type haplogroups in southwestern Hokkaido (37.3%) might be plausible reasons for these regional differences.

This result is consistent with the result of a morphological analysis by Ossenberg et al. (2006). They described that, among theAinu in Hokkaido, individuals in southeastern Hokkaido(this area is contained within our category of “northeastern/central Hokkaido”)are the closest to the Jomon people, whereas theindividuals in western Hokkaido(这个区域包含在我们的类别的“苏thwestern Hokkaido”)are the closest to mainland Japanese.This result is considered reasonable, given the geographical proximity of southwestern Hokkaido to the main island of Japan.

***However, surprisingly, there wereno regional differences in the frequencies of the Okhotsk‐type haplogroups在北北海道西南部(35.3%和34.9%eastern/central Hokkaido). This indicates that the genetic influence of the Okhotsk culture people diffused rapidly in the fledgling Ainu.

As described earlier, Segawa (2007) stated that the invasion of the Satsumon culture people into the areas inhabited by the Okhotsk culture people and the subsequent decline of Okhotsk culture occurred rapidly.The Okhotsk culture people were considered to have been assimilated rapidly into the Satsumon culture during this process, and became part of the basis of the Ainu.

The Native American connection

Current molecular genetic evidencesuggest that the initial founders of the Americas emerged from an ancestral population of less than 5,000 individuals that evolved in isolation, likely in Beringia, from where they dispersed south after approximately 17 kya.Recent findings about the peopling of northern Asia reconstructed by archaeologists suggest that modern humans (haplogroups C and D) colonized the southern part of Siberia around 40 thousand years ago (kya, pre-LGM) and the far northern parts of Siberia and ancient Beringia, by approximately 30 kya.

Haplogroup D4, the most represented of D clades, is subdivided into fifteen principal subclades (D4a–D4j, D4k’o’p’, D4l–D4n, D4q), which range from ~6 to ~28 kya when using the sequence variation of the entire genome and from ~3 to ~42 kya but all subclades are found in eastern Asia, with the eastern Asian lineages the oldest among them.

The ancient mtDNA D4 have been found in the vicinity of Devil’s Gate, i.e. Northeast Asia, the2017 studyof 7700 bp neolithic hunter-gatherers from Devil’s Gate concluded D4 continuity through to today.:

“The mitochondrial genome of the individual with higher coverage (DevilsGate1) could be assigned to haplogroup D4; this haplogroup is found in present-day populations in East Asia (11) and has also been found in Jomon skeletons in northern Japan …

Modern populations that live in the same geographic region as Devil’s Gate have the highest genetic affinity to our ancient genomes(Fig. 2), with a progressive decline in affinity with increasing geographic distance (r2 = 0.756, F1,96 = 301, P < 0.001; Fig. 3), in agreement with neutral drift leading to a simple isolation-by-distance pattern. TheUlchi, traditionally fishermen who live geographically very close to Devil’s Gate and are the only Tungusic-speaking population from the Amur Basin sampled in Russia (all other Tungusic speakers in our panel are from China), are genetically the most similar populationin our panel. Other populations that show high affinity to Devil’s Gate are the Oroqen and the Hezhen—both of whom, like the Ulchi, are Tungusic speakers from the Amur Basin—as well as modern Koreans and Japanese. Given their geographic distance from Devil’s Gate (Fig. 3), Amerindian populations are unusually genetically close to samples from this site, in agreement with their previously reported relationship to Siberian and other north Asian populations.”

mtDNA haplogroup D is found frequently throughout East Asia, in higher frequencies in Northeast Asia, Siberia and Central Asia. A previous view is Derenko’s who is in favour of an East Asian homeland:

A2010 Russian Derenko studyclarified that D4 has an age of 24–28 kya and is further subdivided into fifteen principal subclades (D4a–D4j, D4k’o’p’, D4l–D4n, D4q), which range from ~6 to ~28 kya. These D4 subclades have a very distinctive geographic distribution, which is highly informative about the demographic history of thenorthern Asia. The study showed that all the subclades are found in eastern Asia and that they had already expanded before the LGM, with their oldest lineages being present in the eastern Asia. In particular,most of the eastern Asian subclades of haplogroup D show coalescence ages of between 15 and 42 kya, thus suggesting that some of them were already present here before the LGM.

The mitochondrial DNA gene marker (Y-haplogroup C-M217*) are genetic characteristics also shared by the Nivkhi in northern Sakhalin, and Koryaks in the Kamchatka Peninsula. Hence, the Ainu can be said to be related to the Nivkhi and the Koryaks. However, since the Nivkhi do not carry haplogroup D which has a dominant presence in the Ainu, the migration is seen to have occurred unidirectionally, from the North into Hokkaido. This appears to be confirmed by theresearch of the Saitou laboratory at the National Institute of Genetics, which concluded that the Ainu of Hokkaido (as well as ancient Jomon) showed the closest genetic affinity to native Okinawans.

Another2020 study on the neolithic Boisman cultureobserved continuity going back 8,000 years of the D4 ancestry in the Amur River Basin, as well as its affinity with the Jomon:

“The individuals from the ~5000 BCE Neolithic Boisman culture and the ~1000 BCE Iron Age Yankovsky culture together with the previously published ~6000 BCE data from Devil’s Gate cave19 are genetically very similar, documenting a continuous presence of this ancestry profile in the Amur River Basin stretching back at least to eight thousand years ago (Figure 2 and Figure S2). The genetic continuity is also evident in the prevailing Y chromosomal haplogroup C2b-F1396 and mitochondrial haplogroups D4 and C5 of the Boisman individuals, which are predominant lineages in present-day Tungusic, Mongolic, and some Turkic-speakers. TheNeolithic Boisman individuals shared an affinity with Jomon as suggested by their intermediate positions between Mongolia_East_N and Jomon in the PCAand confirmed by the significantly positive statistic f4 (Mongolia_East_N, Boisman; Mbuti, Jomon).

The expansion of haplogroup D4 settlerstook place in northern Asia post LGM.Of the subclades shared with eastern Asians, D4b1a is thought to represent aseparate Upper Paleolithic migration initiated northward from the Altai-Sayan region of southern Siberia around11-20 kya. D4b1a分为两个分支,其中一个,D4b1a2,largely restricted to northern Asia and its major subclade, D4b1a2a, resulted from the earliest split from the Yukaghir mtDNA within D4b1a2.”

The Paleolithic Siberian population expansions are also thought to be responsible for a migration to the Americas and that North America may have had ancestral relatives of the Jomon and Ainu among its early settlers. SeeAnalysis of HLA genes and haplotypes in Ainu (from Hokkaido, northern Japan supports the premise that they descend from Upper Paleolithic populations of East Asia,Tissue Antigens.2000 Feb;55(2):128-39:

The Ainu people are assumed to be the descendants of pre-agricultural native populations of northern Japan, while the majority of population of present-day Japan (Hondo-Japanese) is considered to have descended mainly from post-neolithic migrants. Sequence-level polymorphisms of the HLA-class I (HLA-A and HLA-B) genes were investigated in DNA samples of 50 Ainu living in Hidaka district, Hokkaido.HLA-A*2402, A*0201, A*0206, A*2601, A*3101, B*1501, B*5101, B*3901, and B*3501 were observed at frequencies of more than 10% and most of these have previously been found in populations of not only Asians but also North and South American Indians.A*68012, which has not so far been detected in Hondo-Japanese, was found in the Ainu (3%).

Correspondence and neighbor-joining analyses of various populations based on HLA-A, -B and -DRB1 gene frequencies enabled distinction between Asian, Native South American, European, and African populations. The Ainu, as well as Tlingit (Na-Dene), were placed midway between other East Asians, including Hondo Japanese, and Native South Americans (Amerindians) in the correspondence analysis. Furthermore,several HLA-A-B and HLA-B-DR-DQ haplotypes common in the Ainu, are shared with some Native American populations. These observations strongly suggest a unique place for the Ainu as descendants of some Upper Paleolithic populations of East Asia, from whom some Native Americans may have descended.

The Ainu are thought to be connected to ancestry that might be in common with those of Native Americans – one connection lies in the detection of the common presence ofHaplogroup Xin both populations. Haplogroup X is traceable to ancient remains of Altaians in the Gobi desert, as well as to the Altai populations today, but ultimately tothe Druze, Basque, and other European populations of the Caucasus. Although it is not at the moment, known how the Haplogroup X arrived in the Ainu ancestral lines, nor to Native American populations in ancient times, a1998 studyconcluded that Native American founders may have had Caucasian ancestry. For more on this seeDNA analyses and inferred genetic origins of the Ainu.

On the other hand…

Astudy on the HLA gene (alleles and haplotypes) frequencieson North, Meso, South American AmeriIndians showed that AmerIndians have little relatedness with Asians, according to genealogy studies except thatNorth-Americans only share one haplotype (A*24-B*40-DRB1*1401-DQB1*0503) with Taiwanese and Japanese in low frequencies.

More work should be done in interdisciplinary studies on virus and disease markers to study ancient migrations. Early work in onestudyshows that all type 2A JC Virus strains from North and South Americans are closely related to strains in present-day Japan. TheJC病毒株出现在纳瓦霍在新墨西哥o (speakers of an Athapaskan language in the Na-Dene language phylum) were found to be of a prototype type 2A strain of a northeast Asian genotype found in Japan. The partialVP1 gene sequences of the JC virus from the Salish people in Montana (speakers of a language in the Salishan group in the Amerind family) and from the Guarani Indians of Argentina (speakers of the Tupi-Guarani language and Equatorial branch of the Amerind family) were however shown to be closely related to several strains variants of strains found in Japan.


See also

References and further reading:

Read more atDNA analyses and inferred genetic origins of the Ainu

Adachi N, Kakuda T, Takahashi R, Kanzawa-Kiriyama H, Shinoda KI.Ethnic derivation of the Ainu inferred from ancient mitochondrial DNA data.Am J Phys Anthropol. 2018;165(1):139-148. doi:10.1002/ajpa.23338

To find out more about the Ainu people, watch these video presentations “Ainu: Spirit of a Northern People” by the Arctic Studies Centerand “Ainu: the first peoples of Japan“, a documentary film by Rhawan Joseph, Ph. D


Morphological affinities between Jomon and Ainu: reassessment based on nonmetric cranial traits”: Shigematsu, Masahito; et al. Anthropological Science Vol. 112: 161-172 (2004) doi:10.1537/ase.00092.

Retr. source:Japan Science and Technology Information Aggregator, Electronic

Origins and genetic features of the Okhotsk people, revealed by ancient mitochondrial DNA analysis”. Sato, Takehiro; et al. (2007). Journal of Human Genetics 52 (7): 618–627. doi:10.1007/s10038-007-0164-z

Mitochondrial DNA haplogrouping of the Okhotsk people based on analysis of ancient DNA: an intermediate of gene flow from the continental Sakhalin people to the Ainu

The History and Geography of Human Genesp253, Cavalli-Sforza ISBN 0-691-08750-4

Jomon genes: using DNA, researchers probe the genetic origins of modern Japanese– Science News, Feb 15, 1997 by John Travis

Kennewick Man Skeletal Find May Revolutionalize Continent’s History, Science Daily

ANTHROPOLOGY: Kennewick Man’s Contemporaries, Science

The Boone Collection – Image Gallery: Ainu Artifacts. Retrieved on 2008-05-08

Genetic origins of the Ainu inferred from combined DNA analyses of maternal and paternal lineagesSource: Journal of Human Genetics, Volume 49, Number 4, April 2004 , pp. 187-193(7)

NOVA Online – Island of the Spirits – Origins of the Ainu” by Gary Crawford

Origins of the Ainu People of Northern Japan“, Suite 101, Jan 9, 2010 by Paula I. Nielson

Diet officially declares Ainu indigenous’, Japan Times, 7 June, 2008 Ito, M, Am J Hum Genet. 1995 April; 56(4): 951–962.

Y chromosomal DNA variation and the peopling of Japan. M F Hammer and S Horai Am J Hum Genet. 1995 April; 56(4): 951–962 PMC1801189

Evidence for a Possible Asian Origin of YAP+ Y Chromosomes.Am. J. Hum. Genet. 61:462-466, 1997, Tasha K. Altheide and Michael F. Hammer

Chandrasekar A, Saheb SY, Gangopadyaya P, et al. (2007). “YAP insertion signature in South Asia“. Ann. Hum. Biol. 34 (5): 582–6. Doi:10.1080/03014460701556262. PMID 17786594.

Underhill et al (2000). “The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations“. Ann. Hum. Genet. 65 (Pt 1): 43–62.

Claudio M. Bravi et al.,Tracing the origin and geographic distribution of an ancestral form of the modern human Y chromosome, Revista Chilena de Historia Natural, 74: 139-149; 2001

Ongoing Adaptive Evolution of ASPM, a Brain Size Determinant in Homo sapiens, Nitzan Mekel-Bobrov, et al., Science 9 September 2005: 309 (5741), 1720-1722. [DOI:10.1126/science.1116815]

Sjöberg, Katarina V. (1993). The Return of the Ainu: Cultural Mobilization and the Practice of Ethnicity in Japan. Studies in Anthropology and History. 9. Chur: Harwood Academic Publ.. ISBN3718654016. OCLC 27684176.

Tajima, Atsushi; et al. (2004). “Genetic origins of the Ainu inferred from combined DNA analyses of maternal and paternal lineages”. Journal of Human Genetics 49 (4): 187–193.doi:10.1007/s10038-004-0131-x.

Hammer, Michael F.; et al. (2006). “Dual origins of the Japanese: Common ground for hunter-gatherer and farmer Y chromosomes”. Journal of Human Genetics 51 (1): 47–58.doi:10.1007/s10038-005-0322-0.

The Peopling of the Americas: A Complex Issue AmeriIndian, Na-Dene, Aleut and Eskimo first inhabitantsInternational Journal of Modern Anthropology, Int. J. Mod. Anthrop. 3 (2010) Available online athttp://www.ata.org.tn

mtDNA haplogroup X: an ancient link between Europe/Western Asia?American Journal of Human Genetics, 1998 ISSN: 0002-9297

The Emerging Limbs and Twigs of the mtDNA Tree, Mol Biol Evol (2002) 19 (10): 1737-1751

Origin and Post-Glacial Dispersal of Mitochondrial DNA Haplogroups C and D in Northern AsiaDerenko M, Malyarchuk B, Grzybowski T, Denisova G, Rogalla U, et al. 2010 Origin and Post-Glacial Dispersal of Mitochondrial DNA Haplogroups C and D in Northern Asia. PLoS ONE 5(12): e15214. doi:10.1371/journal.pone.0015214

Strains of JC virus in Amerind-speakers of North America (Salish) and South America (Guarani), Na-Dene-speakers of New Mexico (Navajo), and modern Japanese suggest links through an ancestral Asian population.Am J Phys Anthropol. 2002 Jun;118(2):154-68.

Archaeology in the Kuril Islands: Advances in the Study of Human Paleogeography and Northwest Pacific Prehistory, Ben Fitzhug et al., Arctic Anthropology, Vol 39, nos 1-2 pp 69-94, 2002 The article examines the human occupation patterns in the Kuril Islands and touches upon the relationship between the Okhotsk, Satsumon and Ainu in the chain of islands.

From Asia to the Americas by boat? Paleogeography, paleoecology and stemmed points of the northwest PacificShell Midden Research: An Interdisciplinary Agenda for the Quaternary and Social Science, Quaternary International, Vol. 239, Issues 1-2, 1 July 2011, Pages 28-37

Abstract: “Within this vast region, scattered Late Pleistocene technological assemblages that include leaf-shaped bifaces and stemmed projectile points found in coastal or peri-coastal sites from Japan and Kamchatka to western North America, and much of South America may support the idea that a coastal migration contributed to the peopling of the Americas.”

Hong Shi, et al.,Y chromosome evidence of earliest modern human settlement in East Asia and multiple origins of Tibetan and Japanese populationsBMC Biology 2008, 6:45 doi:10.1186/1741-7007-6-45

Genetic, Linguistic and Archaeological Perspectives on Human Diversity in Southeast Asia“, Recent Advances in Human Biology Vol. 8 by Li Jin Mark Seielstad, Chunjie Xiao p. 50

(2019)New Results of Research into the Tokarev Culture of Northern Sea of Okhotsk RegionProblems of Archaeology, Ethnography, Anthropology of Siberia and Neighboring Territories (2019) Vol XXV, DOI – 10.17746/2658-6193.2019.25.432-439, Lebedintsev, Alexander; Grebenyuk, Pavel; Fedorchenko, Alexander; Malyarchuk, Boris; Selin, Dmitry

Sato et al., (2007)Origins and genetic features of the Okhotsk people, revealed by ancient mitochondrial DNA analysis, Journal of Human Genetics 52(7):618-27

Hanihara T, Yoshida K, Ishida H.Craniometric variation of the Ainu: an assessment of differential gene flow from northeast Asia into northern Japan, Hokkaido. Am J Phys Anthropol. 2008;137(3):283-293. doi:10.1002/ajpa.20869

Jeong & Wilkin et al.,Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe, PNAS published ahead of print November 5, 2018https://doi.org/10.1073/pnas.1813608115

Dryomov, S.V., Starikovskaya, E.B., Nazhmidenova, A.M. et al.Genetic legacy of cultures indigenous to the Northeast Asian coast in mitochondrial genomes of nearly extinct maritime tribes. BMC Evol Biol 20, 83 (2020).https://doi.org/10.1186/s12862-020-01652-1

Wang et al.,The Genomic Formation of Human Populations in East Asia

29 responses to “Who are the Ainu people?

  1. Yayoi were Koreans. Jomon were Northern Altaic Tribe.

    • ah this is the summary. thanks!

      • From “Ancient genomics reveals tripartite origins of Japanese populations” the paper concludes that Yayoi are of south of Liaodong i.e. northeast Asian transeurasiatic-tungusic ancestry while the Kofun are of East Asian Han but more admixed ancestry between Yellow River, South Korea and Japanese:
        “we find a clear genetic distinction between the groups of people who arrived to the archipelago during
        the subsequent agrarian and state formation phases of Japanese pre- and protohistory. Genetic data from the Yayoi individuals document the presence of Northeast Asian ancestry in the archipelago as supported from morphological studies (5, 49), while we observe widespread East Asian ancestry in the Kofun. The ancestors characterizing each of the Jomon, Yayoi, and Kofun cultures made a significant contribution to the formation of Japanese populations today. …
        Excess affinity to the Yayoi is observable in the individuals who are genetically close to ancient Amur River populations or present-day Tunguisic-speaking populations (Fig. 4 and fig. S17). Our findings imply that wet rice farming was introduced to the archipelago by people who lived somewhere around the Liaodong Peninsula but who derive a major component of their ancestry from populations further north, although the spread of rice agriculture originated south of the West Liao River basin (55).
        The most noticeable archaeological characteristic of Kofun culture is the custom of burying the elite in keyhole-shaped mounds, the size of which reflect hierarchical rank and political power (1).
        The three Kofun individuals sequenced in this study were not buried in those tumuli (see note S1), which suggests that they were lower-ranking people. Their genomes document the arrival of people with majority East Asian ancestry to Japan and their admixture with the Yayoi population… Han, who have multiple ancestral components. A recent study has reported that people became morphologically homogeneous in the continent from the Neolithic onward (56), which implies that migrants during the Kofun period were already highly admixed …” Best to read the paper in its entiretyhttps://www.science.org/doi/pdf/10.1126/sciadv.abh2419

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  3. Eurasian words for ELEPHANT = Very similar! Ask for Whole Paper.
    Chinese S ee AH N g
    Hakka (south China) S I O N g
    Tocharian A ON K A L A M
    Tocharian B ON K O L M O
    Latvian Z I L O N us
    Saami/Lapp S L O NN
    Tibetan G L A N
    Slavic (MANY languages!) S L O N
    Polish (Slavic variant) S “u” O N
    Vietnamese C O N VOI
    Mongol Z A A N
    Japanese Z O S A N
    AINU ? ? ? ? ?

  4. Interesting page. Good job.

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  6. Hello, I found your blog actually by accident. I was doing some research for my blog and stumbled across yours. I saw how much research that you’ve done and realized I didn’t need to do any further study on my own. Great post and research. I hope you don’t mind if I do a pingback to your blog. If you are curious come and take a look at what I’ve done at hague6185.wordpress.com Thanks.

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  8. Japanese consist of Jomon and Yayoi. Jomon are deep set eyed native Japanese like Ken Hirai. Yayoi are slant eyed Chinese and Korean immigrants like average Japanese. Ainu are the descendant of Jomon in Hokkaido. But Hokkaido doesn’t have so many jobs that Ainu come to Tokyo, Osaka etc to work. If you meet non-Asian looking Japanese, they must be Jomon or Ainu Japanese. I give you a link to tell a difference easily between Jomon and Yayoi.

  9. Iranians historian said that 1500 years ago a group of persian had migrated to the east,china and japan maybe.beacuse arabs and muslems invaded to Iran.maybe Ainus were related to them

  10. I think Ainu people are much closer to Assamese and NE Indians. A book published in 2008 “TONKORI” explains thier cultural and linguistic similarities. Also, the Assamese language has about hundreds of words similar to Ainu languages. Eg:-
    “kara” in Ainu means to do
    “kora” in Assamese also means the same.
    “nei” of Ainu has similar uses of “xei” in Assamese.
    “Meko” means cat in ainu whereas “Meko-ri” means the same in Assamese.
    The musical instruments used by Ainus are also similar to some Assamese instruments like “Tonkori” which is “Tokori” in Assamese.Sword called “Emush” is similar to Assamese/Ahomese sword”Heng-Dang” There are also words which are similar to those used in Tai-Ahom(Extinct language), Bishparia, Khasi, Sanskrit, Khampa, etc.

    • From mtDNA data, the shared genepool appears to more between East Asian components of Assamese (A, F, D, M7, M8, M9, and B (24.0%)) as well as East Asian origined genes of the Japanese population, rather than with Ainu population (D, G, M7a, N9b, Y and very small percentages of B, F, A and M7bc) except for M7 – see Shinodahttp://jspsusa.org/FORUM2012/presentation/3-2_Shinoda.pdf
      SeeMeasuring mitochondrial DNA diversity and demographic patterns of tribal and caste populations from the Northeast Indian State of Assam,by PETER H. REJ1, RANJAN DEKA3 and HEATHER L. NORTON2. April 11, 2014
      The Northeast Indian state of Assam has been under-represented in studies of Indian mitochondrial (mtDNA) diversity. To address this gap, sequence data from the HVR-1 and HVR-2 regions and select diagnostic mutations were used to characterize patterns of intra- and inter-population diversity in 205 individuals from tribal (Ahom, Sonowal Kachari, and Rabha) and caste (Assamese) populations of the region. Similar to reports by other studies conducted in India and Southeast Asia, an AMOVA result of 97% variation observed within Assam’s populations demonstrated the heterogeneity of the region.Results indicate that the people of Assam predominantly express autochthonous Indian haplogroups: M, M2, M3, M4, M5, M6, R, R5 (52.5%), but also express a higher degree of East Asian markers: A, F, D, M7, M8, M9, and B (24.0%), due to their close proximity to South China and Southeast Asia. A general lack of Northern Chinese, Tai, and Indo-European ancestral markers, however, indicates that many of the population migrations to Assam were male specific; the ethnically Tai Ahom in particular reflect next to no Southeast Asian specific haplotypes (7.7%), indicative of their 13th century male specific migration. Haplogroup affiliation was more closely associated with geographical location rather than ethnolinguistic heritage. Principal component analysis showed distinct clustering with other Indian caste and tribal populations. This study also identified two novel haplogroups: one subhaplogroup was found within macrohaplogroup M, the other in haplogroup R. Both novel haplogroups were discovered in the Rabha, an ethnically Tibeto-Burmese tribe hailing from Lower Assam. The study concluded:In general, Austro-Asiatic speakers tend to lack haplogroups A, F, D, M7, M8, and B, all of which are common East Asian markers. However, Y markers observed in other studies tend to be Southeast Asian in origin. Based on this sex-biased admixture, one can infer that the founding local female population was present prior to an initial, male-specific, Southeast Asian diaspora that potentially brought the Austro-Asiatic linguistic family from the region, replacing a now extinct language (Metspalu, et al., 2004; Thangaraj et al., 2005).
      When compared to other Caste populations from Metspalu’s 2004 study, the Assamese castes exhibit far more mtDNA markers indicative of East Asian heritage (Metspalu et al., 2004). The general upward social mobility of females, and to a lesser extent the Ahom invasion, resulted in the presence of tribal/East Asian markers in Assamese samples…
      This result accords with another earlier studyhttp://www.ncbi.nlm.nih.gov/pmc/articles/PMC1204953/pdf/ge1311191.pdfwhich found common ancestral components between Assamese, Cambodians and Chinese populations.
      There could be shared genetic components between Assamese and Ainu, perhaps based in Y-DNA analysis, but I don’t have this data on Assamese as yet. The tonkori is an ancient instrument and existed amongst most ancient peoples in E. Asia, SEA but is found throughout Central Asia and Eurasia as well, probably proliferated throughout the Silk Road or even earlier, so its presence is so widespread that it’s hardly an indication of common origin.

      • Ainu and Jomon Japanese people Autosomal Regional DNA were more closely related with Nivks, Udegey and Eastern Siberian DNA or simply a Northeast Asian Typicall Autosomal DNA doesn’t it? A lot of Netizen seems confuse about these because almost all of Ainuid and, perhaps, Jomon People have wet Earwax, Sundadonty dental shapes and more hairy body but Japanese Yayoi generally have dry Earwax and Sinodont dental shapes (EDAR370a).

  11. I’ve just met a Aniu woman whom speaks 5 languages. Most interesting
    I’ve never heard of these people before.

    • Hello, my daughter is doing a year 12 project on Ainu people but it needs input with today’s perspective of Ainu people, so I am looking for a contact. Can you please put us in touch with this woman? She sounds perfect for an interview via email or Facebook etc. If anyone can help at all, please email..gavintina@bigpond,com Thank you, Tina

  12. @Adrian yes, you’re right. The confusion is due to the later admixtures from the Amur, Sakhalin islands and Siberia with indigenous locals from Japan. Shinoda’s studies examining all the ancient dna from Jomon in each region are the clearest efforts to separate out the different lineages and arrivals in terms of time and distance. His graphs and charts show clearly which genetic lineages were present, and which expanded through the ages. We have to be careful with concluding origins based on dry and wet wax though. The common original idea that dry wax is from the north and wet is from the south, is simplistic and no longer holds water. Instead, the dry wax gene is a mutation that developed in the north to northeast Asian regions, long after the Asian lineage had already diverged or separated from the European lineages…is the most recent analysis. Same with early morphological studies that decided that Jomon/Ainu teeth and cranium resembled those of Mungo or Java man and came from Sundaland were conclusions made during an era when much fewer or ancient DNA were available. Today’s more recent morphological analyses show Jomon teeth (size and shape) and cranium are on a cline in between European, Jomon, Mongoloids and Native Americans. Thus, these later analyses are a better match with HLA and virus marker studies.

  13. Kenniwick man who some scientists thought might be related to the Ainu, turned out to be Haplogroup X2a. He was closest genetically related and ancestral to only Native Americans, all of them. As for the Ainu, they have zero instances of haplogroup X2a, X2b or any haplogroup X. The Ainu are most closely related to modern Japanese and other East Asian. They have no ANE or or West Eurasian, unlike the Native Americans. Native Americans are very unique in their ancestry. The Ainu are basal to the Japanese but both are East Asians.

  14. Hope I’m not stepping on any toes here. I was born in Northern Japan and believe I am part Ainu (my father is American). My mother never spoke of her heritage when I inquired. It was like an unspoken subject. Have been looking for a place to get my DNA tested for Ainu heritage more specifically. Unable to find. Can anyone out there help me? And thank you for this website. It’s addictive.

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  23. 你好,我已经为35维罗纳的官方指南years meeting people from many different countries. In particular ( as I have studied some ancient and modern languages and the History of the Arts ) I collaborated with a friend of mine Manager of the Contemporary Art Gallery in organizing an INUIT ART Exibition. A new world opened to my eyes! I spoke to some groups of Inuits coming to the Exibition ( a marketing operation to let the Italians understand that Culture) and met 2 “SHAMAN” together with some hunters and fishermen, some sculptors and some lady-singers producing guttural songs…similar to the songs of the 3 AINU Official Singers I saw in a TV documentary….same melody and sound !!! Shall I understand that the INUIT have a same origin of the AINU ? Is there anyone who can give me an answer? Thank you so much!
    Prof. Mauro Albrigi

  24. Ingvald Karlsson

    Thank you so much for your very fine work on the ainu-people. I live in Sweden and our indigenous people are called “Samer”.

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