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Jōmon people

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Diorama of Jomon people at Sannai Maruyama

Jōmon people (縄文 人, Jōmon jin) is the generic name of the indigenous hunter-gatherer population that lived in the Japanese archipelago during the Jōmon period (c. 14,000 to 300 BC). They were united through a common Jōmon culture, which reached a considerable degree of sedentism and cultural complexity.

The Jōmon people are characterized by a deeply diverged East Asian ancestry and contributed around 10–20% ancestry to modern Japanese people.[1][2][3][4] Population genomic data from multiple Jōmon period remains suggest that they diverged from "Ancestral East Asians" prior to the divergence of Northern and Southern East Asians, sometime between 38,000 and 25,000 years ago, but after the divergence of "Basal East Asian" Tianyuan and Hoabinhian lineages (c. 39,000 years ago). After their migration into the Japanese archipelago, they became largely isolated from outside geneflow at c. 20,000 to 25,000 BC.[4][2][3][5]

Culture

[edit]
See also: Jōmon period

The culture of the Jōmon people was largely based on food collection and hunting, but it is also suggested that the Jōmon people practiced early agriculture.[6] They gathered tree nuts and shellfish, were involved in hunting and fishing, and also practiced some degree of agriculture, such as the cultivation of the adzuki bean and soybean.[7] The Jōmon people also used stoneware and pottery, and generally lived in pit dwellings.[8]

Some elements of modern Japanese culture may have come from the Jōmon culture. Among these elements are the precursory beliefs to modern Shinto, some marriage customs, some architectural styles, and possibly some technological developments such as lacquerware, laminated yumi, metalworking, and glass making.

Pottery

[edit]

The style of pottery created by the Jōmon people is identifiable for its "cord-marked" patterns, hence the name "Jōmon" (縄文, "straw rope pattern"). The pottery styles characteristic of the first phases of Jōmon culture used decoration created by impressing cords into the surface of wet clay, and are generally accepted to be among the oldest forms of pottery in East Asia and the world.[9] Next to clay pots and vessels, the Jōmon also made many highly stylized statues (dogū), clay masks, stone batons or rods and swords.[10]

Craftsmanship

[edit]
Magatama – kidney-shaped beads – are commonly found in Jōmon period Japanese finds, as well as in parts of Northeast Asia and Siberia.

There is evidence that the Jōmon people built ships out of large trees and used them for fishing and traveling; however, there is no agreement as to whether they used sails or paddles.[11] The Jōmon people also used obsidian, jade and different kinds of wood.[12] The Jōmon people created many jewelry and ornamental items; for instance, magatama were likely invented by one of the Jōmon tribes, and are commonly found throughout Japan and less in Northeast Asia.[10]

Religion

[edit]
Main article: Ko-Shintō

It is suggested that the religion of the Jōmon people was similar to early Shinto (specifically Ko-Shintō). It was largely based on animism, and possibly shamanism. Other similar religions are the Ryukyuan and Ainu religions.[13]

Languages

[edit]

It is not known what language or languages were spoken in Japan during the Jōmon period. Suggested languages are: the Ainu language, Japonic languages, Austronesian languages, or unknown and today extinct languages.[14][15] While the most supported view is to equate the Ainu language with the Jōmon language, this view is not uncontroversial or easily acceptable as there were probably multiple distinct language families spoken by the Jōmon period population of the Japanese archipelago.[16]

Alexander Vovin (1993) argues that the Ainu languages originated in Central Honshu, and were later pushed northwards into Hokkaido, where the early Ainu-speakers merged with local groups, forming the historical Ainu ethnicity. Bilingualism between Ainu and Japanese was common in Tohoku until the 10th century.[17][18] According to Vovin (2021) there is also some evidence for the presence of Austronesian languages close to the Japanese archipelago, which may have contributed some loanwords to the early Japanese.[18]

Some linguists suggest that the Japonic languages may have been already present within the Japanese archipelago and coastal Korea, before the Yayoi period, and can be linked to one of the Jōmon populations of southwestern Japan, rather than the later Yayoi or Kofun period rice-agriculturalists. Japonic-speakers then expanded during the Yayoi period, by assimilating the newcomers, adopting rice-agriculture, and fusing mainland Asian technologies with local traditions.[19]

Origins

[edit]
See also: Genetic history of East Asians

The Jōmon people represent the descendants of the Paleolithic inhabitants of the Japanese archipelago, which became isolated from other mainland Asian groups some 22,000 to 25,000 years ago. They have been described as "one of the most deeply diverged populations in East Asia".[20][21][5][22]

Genetics

[edit]
Phylogenetic position of the Jōmon lineage among other East Eurasians
Demographic history of the Jomon lineage (A) Maximum likelihood phylogenetic tree reconstructed by TreeMix under a model of two migrations
Principal component analysis (PCA) of ancient and present-day individuals from worldwide populations

The Jōmon lineage is inferred to have diverged from Ancient East Asians between 25–38,000 years ago, before the divergence of Ancient Northern East Asians and Ancient Southern East Asians, but after the divergence of the Upper Paleolithic Tianyuan lineage and ancient Southeast Asian hunter-gatherer lineage (Hoabinhians).[23][4][21] Like other East Asian populations, the ancestors of the Jōmon people originated from Southeast Asia and expanded to East Asia via a coastal route. They represent one of the "earliest waves of migration".[24][21][25][26]

The Jōmon lineage displays a closer genetic affinity to Ancient Northern and Southern East Asian lineages, compared to Basal East Asian Tianyuan or Hoabinhian lineages.[21] Beyond their genetic affinity with other Eastern Asian lineages, the Jōmon also display a weak yet marginally significant genetic affinity with the Yana Rhinoceros Horn Site specimen, associated with Ancient North Eurasians (or Ancient North Siberians), which may point to geneflow between both groups prior to their isolation from other East Eurasian populations.[4] This geneflow may be associated with the introduction of microblade technology to northern Japan.[2][27] But northern Jōmon populations like the Ainu also have strong genetic affinities with lowland East Asians and northeast Siberians.[28]

Jomon and Continental Asian contributions to modern Japanese

Full genome studies on multiple Jōmon remains revealed them to carry gene alleles associated with a higher alcohol tolerance, wet earwax, no derived variant of the EDAR gene, and that they likely frequently consumed fatty sea and land animals. They also carried alleles for medium to light skin, dark and fine/thin hair, and brown eyes. Some samples also displayed a higher risk of developing liver spots if spending too much time in the sun.[25][29][2] However, according to a 2016 study, the derived allele of the EDAR gene is found in 25% of Ainu. Also, there are high frequencies of variants of the OCA2 gene and ADH gene cluster among Ainu, which are widespread in East Asia but virtually absent elsewhere.[28]

Genetic data further indicates that the Jōmon peoples were genetically predisposed for short stature, as well as higher triglyceride and blood sugar levels, increasing the risk of obesity. At the same time, it gave them resistance to starvation. Modern Japanese share these alleles with the Jōmon period population, although at lower and variable frequency, inline with the inferred admixture among modern Japanese peoples. Watanabe et. al (2023) states that the genetic predisposition for shorter stature among Japanese people often correlates with high Jōmon ancestry, with the opposite correlating with high continental East Asian ancestry.[25]

Liu et. al (2024) stated that Jōmon admixture in contemporary Japanese people varies depending on region, with admixture being the highest in southern Japan, especially Okinawa (28.5%), followed by northeastern Japan (19%) and western Japan (12%).[30] Previous studies showed that populations from Tohoku, Kanto and Kyushu were more genetically related to Ryukyuan populations whilst populations from Kinki and Shikoku were more related to continental East Asian populations.[25]

Haplogroups

[edit]

It is thought that the haplogroups D-M55 (D1a2a) and C1a1 were frequent among the historical Jōmon period people of Japan. O-M119 is also suggested to have been presented in at least some Jōmon period remains. One 3,800 year old Jōmon man excavated from Rebun Island was found to belong to Haplogroup D1a2b1(D-CTS 220).[31] Today, haplogroup D-M55 is found in about 35%[32] and haplogroup C1a1 in about 6% of modern Japanese people. D-M55 is found regularly only in Japanese (Ainu, Ryukyuans, and Yamato) and, albeit with much lower frequency, in Koreans.[33] D-M55 also has been observed in Micronesia 5.1%, Timor 0.2%, China 0–0.4%, this is explained by recent admixture, dating back to the Japanese empire (1868-1945) occupation of those regions.[33] Haplogroup C1a1 has been found regularly in about 6% of modern Japanese. Elsewhere, it has been observed sporadically in individuals from South Korea, North Korea (South Hwanghae Province), and China (ethnic Korean in Ning'an and Han Chinese in Linghai, Guancheng Hui District, Haigang District, and Dinghai District).[34] A 2021 study estimated that the frequency of the D-M55 clade increased during the late Jōmon period.[35] The divergence between the D1a2-M55 and the D1a-F6251 subclades (the latter of which is common in Tibetans, other Tibeto-Burmese groups, and Altaians, and has a moderate distribution in the rest of East Asia, Southeast Asia, and Central Asia) may have occurred near the Tibetan Plateau.[32]

The MtDNA haplogroup diversity of the Jōmon people is characterized by the presence of haplogroups M7a and N9b. Studies published in 2004 and 2007 show the combined frequency of M7a and N9b observed in modern Japanese to be from 12~15% to 17% in mainstream Japanese.[36][37] N9b is frequently found among the Hokkaido Jomons while M7a is found frequently among the Honshu Jomons.[38] However N9b is found only at very low percentage among the Honshu Jomon.[38] M7a is estimated to share a most recent common ancestor with M7b'c, a clade whose members are found mainly in Japan (including Jōmon people), other parts of East Asia, and Southeast Asia, 33,500 (95% CI 26,300 <-> 42,000) years before present.[39] All extant members of haplogroup M7a are estimated to share a most recent common ancestor 20,500 (95% CI 14,700 <-> 27,800) years before present.[39] Haplogroup M7a now has its highest frequency in Okinawa.

A 2018 study stated that Okhotsk‐type and Siberian-type haplogroups contributed to the genetic makeup of Edo-era Ainu, along with mainland Japanese, contrary to the popular belief that Ainu received little genetic input from mainland Japanese before the Meiji-era colonization of Hokkaido.[40]

Morphological characteristics

[edit]
Male skull of the late Jōmon period (replica). Excavated at Miyano Kaizuka (Iwate Prefecture). Exhibition in National Museum of Nature and Science.[41]

Several studies of numerous Jōmon skeletal remains that were excavated from various locations in the Japanese archipelago allowed researchers to learn more about the Jōmon period population of Japan. The Jōmon people were relatively close to other East Asian people, however shared more similarities with Native American samples. Within Japan, regional variance among different Jōmon remains was detected. Historically, the Jōmon people were classified as Mongoloid.[42][43]

Forensic reconstruction from a Jōmon skull, displayed at Niigata Prefectural Museum of History

Dental morphology suggests that the Jōmon had Sundadont dental structure which is more common among modern Southeast Asians and indigenous Taiwanese, and is ancestral to the Sinodont dental structure commonly found among modern Northeast Asians, suggesting that the Jōmon split from the common "Ancestral East Asians" prior to the formation of modern Northeast Asians.[44]

Kondo et al. 2017, analyzed the regional morphological and craniometric characteristics of the Jōmon period population of Japan, and found that they were morphologically heterogeneous and displayed differences along a Northeast to Southwest cline. Differences were based on the cranial index, with Hokkaido Epi-Jomon crania being mesocephalic and Okinawan crania being brachycephalic. They concluded that the "Jomon skulls, especially in the neurocranium, exhibit a discernible level of northeast-to-southwest geographical cline across the Japanese archipelago, placing the Hokkaido and Okinawa samples at both extreme ends. The following scenarios can be hypothesized with caution: (a) the formation of Jomon population seemed to proceed in eastern or central Japan, not western Japan (Okinawa or Kyushu regions); (b) the Kyushu Jomon could have a small-sized and isolated population history; and (c) the population history of Hokkaido Jomon could have been deeply rooted and/or affected by long-term extrinsic gene flows." They also suggested that regional differences in cranial length is based on genetic, or phylogenetic, and environmental effects.[45] But Gakuhari et al. 2020 clarifies that Hokkaido and Honshu Jōmon cluster together and form a clade to the exclusion of other groups.[2]

According to Chatters et al., the Jōmon display some similarities to the Native American Kennewick Man.[46] Chatters, citing anthropologist C. Loring Brace, classified Jōmon and Polynesians as a single craniofacial "Jōmon -Pacific" cluster.[47] Chatters, citing Powell, argues that the Jōmon most resembled the Native American Kennewick Man and Polynesians. According to him, the Ainu descend from the Jōmon people, an East Asian population with "closest biological affinity with south-east Asians rather than western Eurasian peoples".[46][48] Powell further elaborates that dental analysis showed the Jōmon to be of the Sundadont type.[47] According to Matsumura et. al (2019), the Jōmon shared similar phenotypes with prehistoric south Chinese and Southeast Asian peoples, such as 'dolichocephalic calvaria, large zygomatic bones, remarkably prominent glabellae and superciliary arches, concave nasal roots, and low and wide faces'.[49] According to Wang et. al (2021), these peoples were still related to later East Asians despite phenotypic discontinuities.[50]

Craniofacial features of the Jōmon people were significantly retained by the Ainu and Okinawans/Ryukyuans.[51] The former have 2 genes "associated with facial structure in Europeans" but still possessed hair and teeth morphology found in East Asians.[52] In regards to facial flatness, the Ainu were intermediate between Caucasoids and Mongoloids[53] but another study states that they were well within the Mongoloid range.[54] The latter have a "well-defined and less flat upper face", which is characterized by a prominent glabella and nasal root.[51]

ATL retrovirus

[edit]

A gene common in Jōmon people is a retrovirus of ATL (human T lymphotropic virus, HTVL-I). This virus was discovered as a cause of adult T cell leukemia (ATL), and research was advanced by Takuo Hinuma of Kyoto University Virus Research Institute.

Although it was known that many virus carriers existed in Japan, it was not found at all in neighboring countries of East Asia. Meanwhile, it has been found in many Africans, Native Americans, Tibetans, Siberians, Burmese people, Indigenous people of New Guinea, Polynesians, etc. Looking at distribution in Japan, it is seen particularly frequently in southern Kyushu, Nagasaki Prefecture, Okinawa and among the Ainu. And it is seen at medium frequency in the southern part of Shikoku, southern part of the Kii Peninsula, the Pacific side of the Tōhoku region (Sanriku) and Oki Islands. Overall, carriers of the ATL retrovirus were found to be more common in remote areas and remote islands. When examining the well-developed areas of ATL in each region of Kyushu, Shikoku, and Tōhoku in detail, carriers are preserved at high rates in small settlements that were isolated from the surroundings and inconvenient for traffic.

The path of natural infection of this virus is limited to vertical infection between women and children (most often through breastfeeding) and horizontal infection between males and females (most often from males to females through sexual intercourse).[55]

Based on the above, Hinuma concluded that the high frequency area of this virus indicates that high density remains of Jōmon people.[56]

Contributions to other populations

[edit]

Historical groups

[edit]

Full genome analyses of Okhotsk culture remains on Sakhalin found them to be derived from three major sources, notably Ancient Northeast Asians, Ancient Paleo-Siberians, and Jōmon people of Japan. An admixture analysis revealed them to carry c. 54% Ancient Northeast Asian, c. 22% Ancient Paleo-Siberian, and c. 24% Jōmon ancestries respectively.[57]

Genetic analyses on ancient remains from the southern Korean Peninsula revealed elevated Jōmon ancestry at c. 37%, while Yayoi remains in Japan were found to carry nearly equal amounts of Jōmon ancestry (35–60%) and Ancient Northeast Asian-like ancestry (40–65%). These results suggest the presence of a Jōmon-like population on the Korean peninsula and their significant contribution to the formation of early Japonic-speakers. As such, the "agricultural transition in prehistoric Japan involved the process of assimilation, rather than replacement, with almost equal genetic contributions from the indigenous Jomon" and mainland Asian migrants of the Mumun/Yayoi period.[4][58]

Modern groups

[edit]
The Ainu are among the modern groups displaying the highest amounts Jōmon-derived ancestry.

Jōmon-associated ancestry is commonly found throughout the Japanese archipelago, ranging from c. 15% among modern Japanese people, to c. 30% among Ryukyuan people, and up to c. 75% among modern Ainu people, and at lower frequency among surrounding groups, such as the Nivkhs or Ulch people, but also Koreans and other coastal groups, suggesting that the Jōmon were not completely isolated from other groups.[2][29][57][59][21] Wang and Wang (2022) noted that peoples with Jōmon ancestry during the Three Kingdoms period significantly contributed to the genetic makeup of modern Koreans. But this ancestry was 'diluted' over time due to subsequent arrivals of northern Han Chinese.[58]

McColl et. al (2018) states that present Southeast Asians derive ancestry from at least 4 ancient population sources. One source was the mainland Hòabìnhians, who shared ancestry with the ancient Japanese Ikawazu Jōmon, along with present-day Andamanese Önge and Malaysian Jehai.[60] However, according to Yang et. al (2020), the Jōmon individual showed affinities to several coastal Neolithic populations in Siberia and southern East Asia, with the latter clustering with present-day southern East Asians.[61]

Along with modern Japanese, Ulch, and Nivkhs, present-day Taiwanese aborigines (i.e. Ami and Atayal) also showed excess allele sharing with a 2500 year old individual in mainland Japan, which was characterized by 'typical Jōmon culture'.[2]

[edit]

Aspects of the Jōmon culture and pottery were used in the video game The Legend of Zelda: Breath of the Wild. Nintendo's art director Takizawa Satoru said that the Jōmon culture was the inspiration for the "Sheikah slates, shrines and other ancient objects" in the game.[62]

A recreated Jōmon village in the form of an experience park (Sarashina no Sato), which offers different activities, can be visited in Chikuma, Nagano.[63]

See also

[edit]

References

[edit]
  1. ^ Furuichi, Yu (11 June 2019). "'Jomon woman' helps solve Japan's genetic mystery". NHK WORLD. Archived from the original on 11 June 2019.
  2. ^ a b c d e f g Gakuhari, Takashi; Nakagome, Shigeki; Rasmussen, Simon; Allentoft, Morten E.; Sato, Takehiro; Korneliussen, Thorfinn; Chuinneagáin, Blánaid Ní; Matsumae, Hiromi; Koganebuchi, Kae; Schmidt, Ryan; Mizushima, Souichiro; Kondo, Osamu; Shigehara, Nobuo; Yoneda, Minoru; Kimura, Ryosuke (25 August 2020). "Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations". Communications Biology. 3 (1): 437. doi:10.1038/s42003-020-01162-2. PMC 7447786. PMID 32843717.
  3. ^ a b Osada, Naoki; Kawai, Yosuke (2021). "Exploring models of human migration to the Japanese archipelago using genome-wide genetic data". Anthropological Science. 129 (1): 45–58. doi:10.1537/ase.201215.
  4. ^ a b c d e Cooke, Niall P.; Mattiangeli, Valeria; Cassidy, Lara M.; Okazaki, Kenji; Stokes, Caroline A.; Onbe, Shin; Hatakeyama, Satoshi; Machida, Kenichi; Kasai, Kenji; Tomioka, Naoto; Matsumoto, Akihiko; Ito, Masafumi; Kojima, Yoshitaka; Bradley, Daniel G.; Gakuhari, Takashi; Nakagome, Shigeki (2021). "Ancient genomics reveals tripartite origins of Japanese populations". Science Advances. 7 (38): eabh2419. Bibcode:2021SciA....7.2419C. doi:10.1126/sciadv.abh2419. PMC 8448447. PMID 34533991.
  5. ^ a b Jeong, Gichan; Gill, Haechan; Moon, Hyungmin; Jeong, Choongwon (11 December 2023). "An ancient genome perspective on the dynamic history of the prehistoric Jomon people in and around the Japanese archipelago". Human Population Genetics and Genomics: 1–21. doi:10.47248/hpgg2303040008.
  6. ^ Crawford, Gary W. (October 2011). "Advances in Understanding Early Agriculture in Japan". Current Anthropology. 52 (S4): S331–S345. doi:10.1086/658369.
  7. ^ Takahashi, Yu; Nasu, Hiroo; Nakayama, Seiji; Tomooka, Norihiko (2023). "Domestication of azuki bean and soybean in Japan: From the insight of archeological and molecular evidence". Breeding Science. 73 (2): 117–131. doi:10.1270/jsbbs.22074. PMC 10316305. PMID 37404345.
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  11. ^ 堤隆は旧石器時代の神津島での黒曜石採取については、丸木舟を建造出来るような石器が存在しなかったことから考えて、カヤックのようなスキンボートを使用したのではないかと指摘している（堤隆『黒曜石3万年の旅』NHKブックス、2004年、93ページ）
  12. ^ 本節の典拠は橋口、前掲書、158-172ページ
  13. ^ Pilgrim, Richard; Ellwood, Robert (1985). Japanese Religion (1st ed.). Englewood Cliffs, New Jersey: Prentice Hall Inc. pp. 18–19. ISBN 978-0-13-509282-8.
  14. ^ 小泉保（1998）『縄文語の発見』青土社 (in Japanese)
  15. ^ 『古代に真実を求めて 第七集（古田史学論集）』2004年、古田史学の会（編集） (in Japanese)
  16. ^ "蝦夷とアテルイ". masakawai.suppa.jp (in Japanese). Retrieved 26 March 2019.
  17. ^ Vovin, Alexander (1993). A Reconstruction of Proto-Ainu. BRILL. doi:10.1163/9789004644823. ISBN 978-90-04-09905-0.[page needed]
  18. ^ a b Vovin, Alexander (21 December 2021). "Austronesians in the Northern Waters?". International Journal of Eurasian Linguistics. 3 (2): 272–300. doi:10.1163/25898833-00320006.
  19. ^ Chaubey, Gyaneshwer; Driem, George van (2020). "Munda languages are father tongues, but Japanese and Korean are not". Evolutionary Human Sciences. 2: e19. doi:10.1017/ehs.2020.14. PMC 10427457. PMID 37588351. The Japonic-speaking Early Jōmon people must have been drawn in to avail themselves of the pickings of Yayoi agricultural yields, and the Yayoi may have prospered and succeeded in multiplying their paternal lineages precisely because they managed to accommodate the Jōmon linguistically and in material ways.
    The dual nature of Japanese population structure was advanced by Miller, who proposed that the resident Jōmon population spoke an Altaic language ancestral to modern Japanese, and this Altaic tongue underwent Austronesian influence when the islanders absorbed the bearers of the incursive Yayoi culture.
  20. ^ Adachi, Noboru; Kanzawa-Kiriyama, Hideaki; Nara, Takashi; Kakuda, Tsuneo; Nishida, Iwao; Shinoda, Ken-Ichi (2021). "Ancient genomes from the initial Jomon period: new insights into the genetic history of the Japanese archipelago". Anthropological Science. 129 (1): 13–22. doi:10.1537/ase.2012132. As mentioned above, Jomon people are descendants of a common ancestor, although the process of their formation is still unknown. However, their origin dates back to the Paleolithic period based on the distribution of mitochondrial DNA haplogroups of the Jomon people and the age of divergence, which was 22000–23000 YBP (Adachi et al., 2011), and their phylogenetic basal position in the nuclear genome analysis (Kanzawa-Kiriyama et al., 2019).
  21. ^ a b c d e Yang, Melinda A. (6 January 2022). "A genetic history of migration, diversification, and admixture in Asia". Human Population Genetics and Genomics: 1–32. doi:10.47248/hpgg2202010001.
  22. ^ Cooke, Niall P.; Murray, Madeleine; Cassidy, Lara M.; Mattiangeli, Valeria; Okazaki, Kenji; Kasai, Kenji; Gakuhari, Takashi; Bradley, Daniel G.; Nakagome, Shigeki (June 2024). "Genomic imputation of ancient Asian populations contrasts local adaptation in pre- and post-agricultural Japan". iScience. 27 (6): 110050. doi:10.1016/j.isci.2024.110050. ISSN 2589-0042. PMC 11176660. PMID 38883821.
  23. ^ Osada, Naoki; Kawai, Yosuke (2021). "Exploring models of human migration to the Japanese archipelago using genome-wide genetic data". Anthropological Science. 129 (1): 45–58. doi:10.1537/ase.201215. Most Southeast, East, and Northeast Asian populations, including Jomon, are nearly equally distant from the Tianyuan individual, supporting the hypothesis that the Tianyuan population are diverged from the lineage basal to all East and Northeast Asians.
  24. ^ de Boer, Elisabeth; Yang, Melinda A.; Kawagoe, Aileen; Barnes, Gina L. (2020). "Japan considered from the hypothesis of farmer/language spread". Evolutionary Human Sciences. 2: e13. doi:10.1017/ehs.2020.7. PMC 10427481. PMID 37588377.
  25. ^ a b c d Watanabe, Yusuke; Ohashi, Jun (June 2023). "Modern Japanese ancestry-derived variants reveal the formation process of the current Japanese regional gradations". iScience. 26 (3): 106130. Bibcode:2023iSci...26j6130W. doi:10.1016/j.isci.2023.106130. PMC 9984562. PMID 36879818. Whole-genome analyses extracted from the remains of the Jomon people showed that they were highly differentiated from other East Asians, forming a basal lineage to East and Northeast Asians.8,10,11 The genetic relationship between Jomon individuals and other East Asians suggests that the ancestral population of the Jomon people is one of the earliest wave migrants who might have taken a coastal route from Southeast Asia toward East Asia.11 It was also revealed that the Jomon people are genetically closely related to the Ainu/Ryukyuan population and that 10–20% of the genomic components found in mainland Japanese are derived from the Jomon people.8,10 Recent studies have found that, in addition to the "East Asian" population, which is closely related to modern Han Chinese, the "Northeast Asian" population also contributed to the ancestry of modern Japanese people.12,13 Cooke et al. 202113 showed the deep divergence of the Jomon people from continental populations, including the "East Asians" and "Northeast Asians"; thus, it can be concluded that the modern mainland Japanese are a population with genomic components derived from a basal East Asian lineage (i.e., the Jomon people) and from continental East Asians.
  26. ^ Aoki, Kenichi; Takahata, Naoyuki; Oota, Hiroki; et al. (30 August 2023). "Infectious diseases may have arrested the southward advance of microblades in Upper Palaeolithic East Asia". Proceedings of the Royal Society B: Biological Sciences. 290 (2005). doi:10.1098/rspb.2023.1262. PMC 10465978. PMID 37644833. 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.
  27. ^ Natsuki, Daigo (19 January 2021). "Migration and adaptation of Jomon people during Pleistocene/Holocene transition period in Hokkaido, Japan". Quaternary International. 608–609: 49–64. doi:10.1016/j.quaint.2021.01.009. S2CID 234215606. The Incipient Jomon communities coexisted with the Terminal Upper Paleolithic (TUP) people that had continued to occupy the region since the stage prior to the LG warm period, but the Incipient Jomon population was relatively small.
  28. ^ a b Jeong, Choongwon; Nakagome, Shikegi; Rienzo, Anna Di (2016). "Deep History of East Asian Populations Revealed Through Genetic Analysis of the Ainu". Genetics. 202 (1): 261–272. doi:10.1534/genetics.115.178673. PMC 4701090. PMID 26500257.
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