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2015 paper concludes that migrants from Northeast China via Korea bearing the CY JC virus genotype brought the ALDH2 mutation to Central Japan forming the Yayoi populations

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https://static-content.springer.com/image/art%3A10.1186%2Fs13323-015-0031-1/MediaObjects/13323_2015_31_Fig2_HTML.gif

Fig. 2 a Map showing the distribution of the two major JCV genotypes (CY and MY) in Japan. The areas in which CY and MY were found more frequently (i.e., at rates >75 %) are indicated as CY-rich or MY-rich areas, respectively. The area designated as the intermediate area is where the genotypes CY and MY were found at almost identical frequencies. This map was created utilizing data reported by Kitamura et al. [14]. b The geographic distribution of the ALDH2 variant. The orange area indicates where the frequency of the ALDH2 variant is over 24 % of the population. The ALDH2 mutation-rich area is central Japan.

 

Miyamori, Daisuke et al., Tracing Jomon and Yayoi ancestries in Japan using ALDH2 and JC virus genotype distributions Investigative Genetics 2015 6:14 DOI: 10.1186/s13323-015-0031-1 ©  Miyamori et al. 2015

Overall findings of this study were that the ALDH2 variant is significantly higher in the population with the CY genotype JCV (51.5 %) than in the population with the MY genotype (24.2 %) (p < 0.05) in Japan

Abstract

Background

According to the dual structure model, the modern Japanese ethnic population consists of a mixture of the Jomon people, who have existed in Japan since at least the New Stone Age, and the Yayoi people, who migrated to western Japan from China around the year 300 BC. Some reports show that the Yayoi are linked to a mutation of the aldehyde dehydrogenase 2 gene (ALDH2).

Recent viral studies indicate two major groups found in the Japanese population: a group with the CY genotype JC virus (JCV) and a group with the MY genotype JCV. It is unclear whether either genotype of the JC virus is related to the Jomon or Yayoi.

In this study, we attempted to detect JCV genotypes and ALDH2 mutations from the DNA of 247 Japanese urine samples to clarify the relationship between the dual structure model and the JCV genotype through ALDH2 mutation analysis and JCV genotyping.

Findings

The ALDH2 polymorphism among 66 JC virus-positive samples was analyzed, and it was found that the ALDH2 variant is significantly higher in the population with CY genotype JCV (51.5 %) than in the population with the MY genotype (24.2 %) (p < 0.05).

Conclusion

From these findings, it may be inferred that the ALDH2 mutation, which is related to the Yayoi, is related to CY genotype JCV. When the Yayoi migrated to the Japanese archipelago, they brought the ALDH2 mutation as well as the CY genotype JCV.

JC virus ALDH2 mutation Japanese

Introduction

It is believed that the modern Japanese ethnic population consists of a mixture of the Jomon people, who have existed in Japan since at least the New Stone Age, and the Yayoi people, who migrated to western Japan from China via Korea approximately 2000 to 3000 years ago and were responsible for spreading rice cultivation [25] (Fig. 1). There are many studies that describe this dual structure model of the Japanese people based on analysis of elements such as the human Y chromosome, mitochondria, single nucleotide polymorphism (SNP), archeological data and historical records [3, 7, 8, 9, 10, 19, 20, 21, 26, 28]. Sokal and Thompson [26] interpreted the cause of the genetic distribution to have resulted from the selection, admixture and resolving power of a locus that reflects a particular population event. The aldehyde dehydrogenase 2 gene (ALDH2) is one of the examples among studies.

https://static-content.springer.com/image/art%3A10.1186%2Fs13323-015-0031-1/MediaObjects/13323_2015_31_Fig1_HTML.gif

Fig. 1

This map shows the migration of the Yayoi people and the Jomon people. The Jomon people moved to the north and south of Japan after the Yayoi people migrated to central Japan

Previous studies have shown that approximately 2000 to 3000 years ago, a variant of ALDH2 was present within the Chinese population. Citing historical evidence [18], researchers speculated that people with this ALDH2 variant migrated to western Japan 2000 to 3000 years ago. Oota et al. [22] claimed that based on actual DNA evidence, this ALDH2 variant was a young haplotype associated with low levels of genetic diversity. Therefore, it may be possible that this mutation of the ALDH2 gene and the Yayoi migration from China to Japan happened in roughly the same period.

The JC virus (JCV), one of the polyomaviruses, is a useful marker in tracing the dispersal of human populations [30]. This is because once an individual is infected asymptomatically with JCV during childhood [23, 24], the initially infected JCV strain persists in renal tissue for life, and other strains of JCV are unable to infect the already infected individual [2, 13, 29]. There are more than 20 main JCV genotypes that are distributed in geographically distinct domains throughout the world [30]. Two major types of JCV genotypes, CY and MY, are found in the Japanese archipelago [14] (Fig. 2a). The genotype CY is commonly distributed in western Japan, northeast China, and Korea [5, 6] and is not found in other places [27, 32]. Earlier studies suggested that the Chinese might have brought the CY genotype JCV to Japan when they migrated from China [14]. The genotype MY is commonly distributed in eastern Japan and among Native Americans [27, 32].

https://static-content.springer.com/image/art%3A10.1186%2Fs13323-015-0031-1/MediaObjects/13323_2015_31_Fig2_HTML.gif

Fig. 2

a Map showing the distribution of the two major JCV genotypes (CY and MY) in Japan. The areas in which CY and MY were found more frequently (i.e., at rates >75 %) are indicated as CY-rich or MY-rich areas, respectively. The area designated as the intermediate area is where the genotypes CY and MY were found at almost identical frequencies. This map was created utilizing data reported by Kitamura et al. [14]. b The geographic distribution of the ALDH2 variant. The orange area indicates where the frequency of the ALDH2 variant is over 24 % of the population. The ALDH2 mutation-rich area is central Japan. This map was created utilizing data reported by Li et al. [17]

This JCV distribution may support the dual structure model of the Japanese population. However, there have been no studies verifying the correlation between the distribution of the CY genotype JCV and the dual structure model.

Therefore, through the detection of the ALDH2 mutation among JCV-positive Japanese samples, we investigated its correlation with the CY genotype JCV, and we discuss whether the dual structure model is also supported by JCV genotype in this study. …

Results

JCV was detected in 66 samples. Among them, the genotype CY was detected in 33 samples and the genotype MY was detected in 33 samples. Among the 33 CY detected samples, the ALDH2 variant was detected in 17 samples (51.5 %). Among the 33 MY detected samples, the ALDH2 variant was detected in only 8 samples (24.2 %). All of the ALDH2 variants were Glu/Lys heterozygotes. Lys/Lys homozygotes were not detected in the samples. The ALDH2 variant was found more commonly in people who carry the CY genotype JCV than in people who carry the MY genotype JCV (Table 1) (p value 0.04, odds ratio 0.30).

Table 1

Association between the JCV genotype and the ALDH2 variant

MY genotype

CY genotype

ALDH2 variant

8

17

(Glu/Lys heterozygote)

ALDH2 wild type

25

16

(Glu/Glu homozygote)

Total

33

33

Lys/Lys homozygote was not detected in this study.

Discussion

The results of the present study suggest that the ALDH2 variants are more prevalent in people with CY genotype JCV than in people with MY genotype JCV (Table 1).

In an earlier study, the CY genotype JCV was found to be more common in western Japan, and the MY genotype JCV was found to be more common in Eastern Japan [14] (Fig. 2a). The CY genotype JCV was also found in China [5, 6], but the MY genotype JCV was found mainly in the Japanese archipelago and in North and South America [32]. Estimated from the substitution rate of the JCV genome, the MY clade occurred more than approximately 10,000 to 30,000 years ago [31], and the CY clade occurred approximately 10,000 years ago. Therefore, the MY genotype JCV initially occurred in the Japanese archipelago and spread to the Americas, and later, the CY genotype JCV migrated from China to Japan. It is possible that the ALDH2 mutation only occurred within people who carry the CY genotype JCV, explaining why it is uncommon for people who carry the MY genotype JCV to have the ALDH2 mutation. Another study indicates that Native Americans do not have the ALDH2 mutation [4]. Combining this information with the results of Zheng et al. [32], we can speculate that people who carry the MY genotype JCV may not have originally had the ALDH2 mutation. Those findings also support the relation of the CY genotype JCV with the ALDH2 mutation. Therefore, it is inferred that the ALDH2 gene mutation spread into East Asia in the past few thousand years. This may be a good example of a locus subjected to selection, displaying wide distribution, and high frequency with low associated variation, confined to a continental region.

However, there are some people with the ALDH2 mutation and the MY genotype JCV, and some people are without the ALDH2 mutation and with the CY genotype JCV.

It is believed that there are several reasons for this. First, extensive genetic mixing between Yayoi and Jomon is expected to have occurred after the major migration 2000 to 3000 years ago and before the present day, when our samples were collected. Second, the ALDH2 genes from both parents are passed down and combined forming their child’s ALDH2 gene type. As a result, there is a 50 % possibility of inheriting the ALDH2 mutation from a father or mother carrying the gene. However, in the case of the JCV genotype, the possibility of infection with JCV from one’s father or mother is affected by the number of exposures to their urine. As a result, if the infection rate of one JCV genotype is high within a given area, subsequent generations are less likely to be infected by JCV of other genotypes, which in turn becomes less common through natural selection. Minority groups of viral genotypes are excluded by the dominant genotype within a given area during a given period. These differences in the acquisition of infection from a local majority or minority strain of a virus or the inheritance of a mutant or non-mutant version of a gene on the other might be the cause of the differences in distribution of the JCV genotypes and the ALDH2 mutation in Japan (Fig. 2a, b).

After a study reported evidence of greater genetic affinity between Ainu and Ryukyuan people (i.e., people indigenous to Okinawa and a surrounding chain of islands between Japan and Taiwan) than between either group from the Japanese mainland populations [11], it was suggested that the Jomon migrated to the north and south of the Japanese archipelago in response to the Yayoi migration from China to western Japan (Fig. 1). Because the ALDH2 mutation is thought to be common among Yayoi, the distribution of the CY genotype JCV is believed to be the same as the distribution of those people who have the ALDH2 mutation (Fig. 2a, b). However, Kitamura et al. [14] reported that the most common genotype found in Okinawa is CY. This is contradictory to the idea of the Jomon migration to Okinawa and Tohoku (the northeast region of the main island of Japan) with their MY genotype JCV, following the migration of the Yayoi people to the main islands of the Japanese archipelago. According to our model, all specimens from before the Yayoi arrival would lack both the ALDH2 mutation and the CY viral genotype while bearing the MY genotype. Further studies are necessary to examine other aspects of the Yayoi arrival. The potential value of studying ancient DNA in pre-Yayoi era specimens of the Ryukyuans and other populations in Japan using viral genome capture techniques and ALDH2 mutation analysis could help solve the mystery of the apparent rarity of the MY variant JCV in modern Ryukyuan populations [15].

Conclusion

From these findings, it may be inferred that the ALDH2 mutation, which is related to the Yayoi, is related to CY genotype JCV. When the Yayoi migrated to the Japanese archipelago, they brought the ALDH2 mutation as well as the CY genotype JCV.

Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.

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See also another 2015 paper:

Timothy A. Jinam, Hideaki Kanzawa-Kiriyama and Naruya Saitou, Human genetic diversity in the Japanese Archipelago: dual structure and beyond Genes Genet. Syst. (2015) 90, p. 147–152

The paper concludes that “genetic data strongly support the dual-structure model proposed by Hanihara (1991) whereby the Hondo Japanese are the result of admixture between the Jomon and Yayoi ancestral populations (Fig. 2). The indigenous Ainu and Ryukyuan populations retain a genetic identity that most likely traces back to Jomon ancestors, while at the same time show indications of recent admixture with the Hondo Japanese”…whilst noting that “The genetic substructure in the Hondo Japanese also hints at a more complex model of human migrations and interactions than the dual structure model implies”.



Ancient Japan may have been ‘more cosmopolitan’ than thought: researchers

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image

Physics.org, Oct 5, 2016

Ancient Japan may have been far more cosmopolitan than previously thought, archaeologists said Wednesday, pointing to fresh evidence of a Persian official working in the former capital Nara more than 1,000 years ago.
Present-day Iran and Japan were known to have had direct trade links since at least the 7th century, but new testing on a piece of wood—first discovered in the 1960s—suggest broader ties, the researchers said.
Infrared imaging revealed previously unreadable characters on the wood—a standard writing surface in Japan before paper—that named a Persian official living in the country.
The official worked at an academy where government officials were trained, said Akihiro Watanabe, a researcher at the Nara National Research Institute for Cultural Properties.
The official may have been teaching mathematics, Watanabe added, pointing to ancient Iran’s expertise in the subject.
“Although earlier studies have suggested there were exchanges with Persia as early as the 7th century, this is the first time a person as far away as Persia was known to have worked in Japan (during the period),” he said.
“And this suggests Nara was a cosmopolitan city where foreigners were treated equally.”
Nara was the capital of Japan from around 710 AD to around 784 AD before it was moved to Kyoto and later present-day Tokyo. …
Read more at: http://phys.org/news/2016-10-ancient-japan-cosmopolitan-thought_1.html#bjCp


Hoabinhian sites in Asia and relict DNA

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The oldest Hoabinhian site has recently been discovered in Yunnan, China. What implications does this have for the peopling of East Asia?

It has been loosely pointed out that Hoabinhian artefacts have been found in Japan (roughly corresponding to the Jomon), source: Uncovering Southeast Asia’s Past (see Chapter 33) Sandra Bowdler’s map of distribution of H. artefacts in Asia refers.

Wilhelm G. Solheim in his Archaeology and Culture in Southeast Asia: Unraveling the Nusantao cites as evidence the observations of Gerard J. Groot and P. van Stein Callenfels which find early Jomon axes (especially the finds of Kozanji shell mound) to correspond perfectly to Bacsonian axes, including polished stone tools. ‘It is now considered that the “Hoabinhian and Bacsonian are two parallel developments of the same culture”.’

Read the new May 2016 paper detailing the latest find of the earliest Hoanbinhian artifacts in Yunnan (below.)

Xueping Ji, The oldest Hoabinhian technocomplex in Asia (43.5 ka) at Xiaodong rockshelter, Yunnan Province, southwest China Quaternary International, Vol 200 2 May 2016, Pages 166–174  http://dx.doi.org/10.1016/j.quaint.2015.09.080

Abstract

The Hoabinhian is the most representative technocomplex in Southeast Asian prehistory for the later hunter–gatherer period. As a mainland technology based exclusively on seasonal tropical environments, this core-tool culture was previously defined in northern Vietnam in 1932 and characterized originally by its large, flat and long, largely unifacial cobble tools associated with tropical forest fauna. The recent discoveries and dates obtained at Xiaodong rockshelter in Yunnan Province (southwest China) allow us to discuss the origin and the homeland of this singular Asian technocomplex which spread to Southeast Asia during the end of the Late Upper Pleistocene. Here we present the first Chinese Hoabinhian lithic implements in their stratigraphic and chronological context within a rockshelter site, and we address the question of the dispersal of modern humans from South China to Southeast Asia

There are theories that Hoabinhian tool makers were mtDNA R9 or basal mtDNA N descendants (source). Nothing is really conclusive unless ancient DNA can be found confirming the identity of the Hoabinhian tool makers.

About.com characterizes the Hoabinhian culture as follows:

The Hoabinhian Period is the name given to that part of Southeast Asian prehistory from about 13,000 to 3000 BC. Archaeological evidence at sites such as Spirit Cave (Thailand) and Cai Beo(Vietnam) reveal that people lived in caves, at open air sites, or along coastal locations as hunter-gatherers and fishers. Coastal Hoabinhian sites often have large shell middens. Animal bones recovered from Hoabinhian sites include primarily wild pigs and deer. Plant remains from Spirit Cave have included almond, bamboo, and gourd. No evidence for domesticated millet or rice has been found at Hoabinhian sites to date.

Hoabinhian stone tools were made from pebbles, and include ground stone axes, grindstones and a dominant stone flake industry. Bone points and spatulas carved from animal bone are also known. In the later Hoabinhian period (argued to be about 5000 BC) sites are also found with potsherds, impressed with vines, mats, or cords.

Hoabinhian burials are generally flexed or contracted; they are often covered with hematite, a common trait of hunter-gatherers just about everywhere.

In Ancient Southeast Asia, John Norman Miksic and Goh Geok Yian write

The oldest Hoabinhian artifacts have been dated to the Pleistocene well before the domestication of plants and animals, or horticulture, is believed to have emerged: at Tham Lod (dates 32,400 – 12.100 BP uncalibrated) and Tham Khuong in Vietnam (20,000-28,000 BP, uncalibrated) 

Hoabinhian Archaeological Sites

Tögi Ndrawa (Sumatra), Spirit Cave and Banyan Valley Cave(Thailand), Cai Beo (Vietnam), Tam Hang (Laos.

….

Archaeologists discover oldest Hoabinhian site ever in China

by Matt Atherton, Dec 30, 2015

The oldest site of south-east Asian hunter-gatherer culture has been discovered in China. The site, dating back nearly 44,000 years ago, was found in the Yunnan Province at Xiaodong Rockshelter, south-west China, and is the oldest evidence of ‘Hoabinhian culture’ ever discovered.

Hoabinhian culture describes the life of hunter-gatherers in south-east Asia, and was originally believed to be a time period stretching back as far as 29,000 years ago. The new discovery, made by scientists all over the world including China, South Africa and France, suggests that the Hoabinhian age may have begun a lot earlier than thought.

“Xiaodong can be regarded as a typical early Hoabinhian site, the first such site found in China and currently the oldest in Asia as well,” said Ji Xueping, lead author of the study. “The study shows that the Lancang River valley is a possible home for the Hoabinhian culture, and a source of migration for modern humans and the transmission of their culture to south-east Asia.”

Archaeologists researching Hoabinhian sites look for areas with large axe-like tools, used for woodwork in forested areas. Some of these stone artefacts were first collected at the Xiaodong site in 2004.

Since then, further research has been carried out to try and confirm the scientific value of the site. This research was then verified and published in December 2015, in Quaternary International.

Carbon dating was used to verify the date from which the site was occupied. Four metres of vertical soil layer was dated – showing the site was inhabited between 43,500 to 24,000 years ago. The scientists speculated that the very bottom layer may be even older.

The discovery provides further insight into how hominids survived a long time ago. It shows some of their strategies for survival, as well as how – and when – they changed from sporadic settlements to agricultural communities.

Hoabinhian culture was first discovered in Vietnam nearly 100 years ago. Since then, more site discoveries in south-east Asia have been dated back between 29,000 and 5,000 years ago. This is the first of these sites to be found in China. … Read more at IB Times

For a discussion of what DNA might have constituted the oldest arrivals in Southeast Asia, including the Hoabinhian culture, see Hill, Catherine, Phylogeography and Ethnogenesis of Aboriginal Southeast Asians Mol Biol Evol (2006) 23 (12):2480-2491.doi: 10.1093/molbev/msl124:

Abstract

Studying the genetic history of the Orang Asli of Peninsular Malaysia can provide crucial clues to the peopling of Southeast Asia as a whole. We have analyzed mitochondrial DNA (mtDNAs) control-region and coding-region markers in 447 mtDNAs from the region, including 260 Orang Asli, representative of each of the traditional groupings, the Semang, the Senoi, and the Aboriginal Malays, allowing us to test hypotheses about their origins. All of the Orang Asli groups have undergone high levels of genetic drift, but phylogeographic traces nevertheless remain of the ancestry of their maternal lineages. The Semang have a deep ancestry within the Malay Peninsula, dating to the initial settlement from Africa >50,000 years ago. The Senoi appear to be a composite group, with approximately half of the maternal lineages tracing back to the ancestors of the Semang and about half to Indochina. This is in agreement with the suggestion that they represent the descendants of early Austroasiatic speaking agriculturalists, who brought both their language and their technology to the southern part of the peninsula ∼4,000 years ago and coalesced with the indigenous population. The Aboriginal Malays are more diverse, and although they show some connections with island Southeast Asia, as expected, they also harbor haplogroups that are either novel or rare elsewhere. Contrary to expectations, complete mtDNA genome sequences from one of these, R9b, suggest an ancestry in Indochina around the time of the Last Glacial Maximum, followed by an early-Holocene dispersal through the Malay Peninsula into island Southeast Asia.

mtDNA haplogroups M21 and R21 were suggested as ancentral in the Semang and Senoi, F1a1a a Holocene component from Indochina.

 

R9b and perhaps N9a are found in high frequency only in Aboriginal Malays … suggest a Pleistocene origin to the north in indochina, wiht an early Holocene dispersal southwards through the Malay Peninsula and into island Southeast Asia.  echoes … the suggestion of Van Heekeren (1072) that the Hoabinhian had originated in South China before spreading to Malaya and North Sumatra.

[What of Austronesian  expansion in the archipelago?]

… on the other hand, haplogroups N21, N22 and M7c1c suggest an equally large offshore component, dating ot the mid/late Holocene, in the ancestry of Aboriginal Malays.

Perhaps the most striking signal is the presence of F1a1a, which aside from the apparently indigenous R21 is the most common haplogroup in the Senoi[R21 diverged from the common haplogroup R ancestor ~60,000 years ago (Macaulay et al. 2005), although the putative control-region link with haplogroup R9 (at np 16304) may imply a younger common ancestor], carried by almost half of the individuals sampled. This haplogroup, which is of early to mid-Holocene age, has b een observed elsewhere at high frequences only in Indochina and probably dispersed there from South China (where it is less frequent but more diverse and where its 1-step ancestor is found) during the Holocene). This suggests that almost half of the maternal lineages of the Senoi may trace back to an origin in Indochina at some point within the last 7,000 years or so.

Gorman, Chester. 1969. Hoabinhian: A Pebble-Tool Complex with Early Plant Associations in Southeast Asia. Science 163(3868):671-673.

Harvick, Ben. 2006. A Methodological Study of Technological Attributes in Hoabinhian Lithic Assemblages. In Social, Cultural and Environmental Dynamics in the Highlands of Pangmapha, Mae Hong Son Province: Integrated Archaeological Research into the Region Rasmi Shoocongdej, ed. Australian National University.

Hoabinhian, Jomon, Yayoi: Early Korean States (Oxbow Monograph) by Gina Lee Barnes

Mineral Food in the late Palaeolithic Hoabinhian Culture of Vietnam and Geophagia in Today Vietnam”, which was presented in the Symposium for Salt Production held in Tokyo Nov 2009, as a part of the paper “Further Studies on Hoabinhian” presented at the IPPA conference, Hanoi, Dec. 2009 and as a paper “Mineral Resources and Salt using in Prehistory Vietnam in comparison [comparation- sic] with Japanese Prehistory


The people and stories behind Meiji Era’s (1867-1926) architecture

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Facade of the State Guest House (Akasaka Palace)

Facade of the State Guest House (Akasaka Palace). Built in 1909, it is one of the biggest buildings constructed during the Meiji period. Photo: Heritage of Japan

Behind every building there is a backstory of the people for whom and the purpose for which it was built. There is also the story of pioneers at the dawn of Western-style architecture in Japan. The buildings of the Meiji Era tell of a bygone era, of the incredible innovations of Industrial Revolution introduced from Europe into Japan, of the local events and developments that paralleled those in the West. Take a walk back in time to explore some of those stories.

Factories are iconic symbols of the Industrial Revolution era. During the 19th century, the worldwide demand for cloth had grown so much that merchants and nations competed fiercely to meet the supply of textiles for making clothing. Textile mills and factories, beginning with England, installed new machines to increase productivity and to cut the costs of labour.

Tomioka Silk Mill

Tomioka Silk Mill is Japan’s oldest silk-reeling factory. Established in 1972 to mass-produce and export high-quality silk

Tomioka Silk Mill, UNESCO site

Tomioka Silk Mill, UNESCO World Heritage site (Source)

Inside: 300 machines imported from France

Inside Tomioka Silk Mill: 300 machines imported from France (Source)

Ukiyoe print of the Tomioka Mill at work

Ukiyoe print of the Tomioka Mill and workers at work (Source)

The Meiji oligarchy wanted the fruits of Western progress, so they sent learning missions abroad to absorb as much of the western technological innovations as possible. One such mission, led by Iwakura, Kido, and Okubo and containing forty-eight members in total, spent two years (1871-73) touring the United States and Europe, studying government institutions, courts, prison systems, schools, the import-export business, factories, shipyards, glass plants, mines, and other enterprises. [Source: Library of Congress]

The government reorganized itself and also formed a professional corps of diplomats. Consequently, many Western-style buildings were built to receive foreign experts and carry out the reforms. Many of the Meiji era buildings were designed by Dr. Katayama Tokuma, who is regarded as the father of modern architecture in Japan. Katayama studied for in many years in the UK, France and Germany and under British architect Josiah Conder.

Aerial view of the Akasaka palace

Aerial view of the Akasaka palace. Photo: Cabinet Office

Among the more extravagant projects undertaken was the Akasaka Palace (see photo above). Styled in part after the New Palace in Vienna, Buckingham Palace and the Louvre, its basic design was based on the Versailles Palace and it remains the only neo-Baroque European style palace in Japan.

Built of stone aroundd a steel frame, the palace building is both fire-resistant and earthquake-proof

Built of stone aroundd a steel frame, the palace building is both fire-resistant and earthquake-proof Photo: Heritage of Japan

Building began in 1899 and was completed in 1909, under the overall direction of Dr. Katayama Tokuyama who was an architect who studied under British architect Josiah Conders, who advised the Japanese government on many projects. The interiors lavishly decorated with imported French mosaics, fireplaces, furniture and chandeliers, and Italian rose and white marble in the early 19th c. Empire style during the rule of Napoleon Bonaparte, Akasaka Palace is, like its Western counterparts, a stately and enduring monument of great beauty. Take a virtual tour of the palace here.

Akasaka (Togu) Palace designed by Katayama Tokuma in 1899, now functions as the State Guest House

Akasaka (Togu) Palace designed by Katayama Tokuma in 1899, was built for the Crown Prince and his bride. Little used as it was deemed expensive, costly to run, instead it was used to house some government departments and the Diet Library. Recently refurbished, it now functions as the State Guest House. The garden fountain seen here features four griffin statues. (Photo credit: Heritage of Japan)

Inside: Stunning and sumptuous staterooms like this ballroom

Inside are many stunning and sumptuous staterooms like this ballroom. The palace fuses Japanese motifs such as samurai armour and helmets,  Japanese drums, phoenix with western flying horses, goddesses and sphinxes in the gilded stucco reliefs.

As part of the “Japanese spirit, Western knowledge” effort, Japanese began wearing European-style army uniforms, morning coats, top hats and ball gowns. Members of the Imperial court adopted European European titles and rococo, neo-baroque, neo-classical buildings were established and concerts of classical music were being performed.

josiah-conder-portrait-1

In its efforts to modernize and in adopting the cultures and systems of the West, Japan needed experts to design Western-style buildings. Josiah Conder(see photo above left. Photo: Wikimedia Commons), an architect was invited from Britain to fill the role as adviser to the Meiji government. One of Conder’s notable projects include the former Iwasaki Family House and Kyu-Iwasaki-tei Garden at Taito-ku Tokyo Japan, designed in 1896(see photo below, photo: Wikimedia Commons)

1024px-former_iwasaki_family_house_and_garden_2010Katayama-Tokuma.jpg

Conder taught the history and structures of Western architecture to Japanese students. His very first class of graduates included Katayama Tokuma(see photo above right, credit: Wikimedia Commons), the man who became court architect (who built the Crown Prince’s Akasaka Palace – see photos above) and who designed the Hyokeikan (see photo below).

Hyokeikan designed by Katayama (student of Josiah Conder), now part of Tokyo Ueno Museum

Hyokeikan, designed by Katayama Tokuma (student of Josiah Conder) and completed in 1908, it now part of Tokyo Ueno Museum (Source: Hyokeikan website)

The Hyokeikan has a magnificent vault dome in its entranceway

The Hyokeikan has a magnificent vault dome in its entranceway (Source: Hyokeikan website)

The word Diet derives from Latin and was a common name for an assembly in medieval Germany. The Meiji constitution was largely based on the form of constitutional monarchy found in nineteenth century Prussia and the new Diet was modeled partly on the German Reichstag.

19th century Japanese Parliament

In keeping with Japan’s modernization and westernization drive, German architects Wilhelm Böckmann and Hermann Ende invited to Tokyo in 1886 and 1887, respectively, drew up two plans for a Diet building which were similar to other Western legislatures of the era. Böckmann’s initial plan, for example, was a masonry structure with a dome and flanking wings, which would form the core of a large “government ring” south of the Imperial Palace. However, due to public resistance in Japan to Foreign Minister Inoue Kaoru‘s internationalist policies, the first two parliament buildings were built to a more “Japanese” design, with traditional Japanese architectural features.

japanese_diet_hall_design_by_fukuzo_watanabe

Japanese Diet Hall Design by Fukuzo Watanabe (Photo: In the public domain)

Prime Minister Katsura Tarō chaired the commission, which recommended that the new building emulate an Italian Renaissance architectural style. This too received much opposition, so after a public design competition in 1918, and 118 design submissions, the current Diet building was built based on the design by the first prize winner, Watanabe Fukuzo, who produced a design similar to Ende and Böckmann’s.

Ende and Böckmann’s Diet Building was never built, but their other “government ring” designs were used for the Tokyo District Court and Ministry of Justice buildings. Returning diplomatic and study missions from the west also called for domestic reforms that were carried out. Between 1871 and 1873, a series of land and tax laws were enacted as the basis for modern fiscal policy. Private ownership was legalized, deeds were issued, and lands were assessed at fair market value with taxes paid in cash rather than in kind as in pre-Meiji days and at slightly lower rates. [Source: Library of Congress *; factsanddetails.com]

Ministry of Justice's former Administration Building, designed by Herman Ende, Wilhelm Bockman in 1895

Ministry of Justice’s former Administration Building, designed by Herman Ende, Wilhelm Bockman in 1895 (Photo: Wikimedia Commons)

From the onset, the Meiji rulers embraced the concept of a market economy and adopted British and North American forms of free enterprise capitalism. The private sector– in a nation blessed with an abundance of aggressive entrepreneurs– welcomed such change.

Economic reforms included a unified modern currency based on the yen, banking, commercial and tax laws, stock exchanges, and a communications network. Establishment of a modern institutional framework conducive to an advanced capitalist economy took time but was completed by the 1890s. [Source: Library of Congress *]

Yokohama Specie Bank

Yokohama Specie Bank, 1880, which became the Bank of Tokyo’s Yokohama branch, now the Kanagawa Prefectural Museum of History and Culture (Photo: Wikimedia Commons)

As in the West, trade and the industrial revolution in Japan created a demand for a host of banking and financial services. Stocks were traded vigorously, new instruments for market speculation evolved on the back of capitalism. New buildings were erected to facilitate and conduct these financial and banking activities. Education and school institutions had to be reformed as well. Germany was the primary model for educational reform. German language and literature were taught and Western-styles schools for the elite were built.

Tokyo University built in the Uchida Gothic style

Tokyo University built in the Uchida Gothic style  Source: Wikimedia Commons

Educational institutions and western-style university buildings such as Tokyo University (above and below) and Meiji University were established in keeping with the necessary educational reforms.

Tokyo University

Tokyo University  Photo: Wikimedia Commons

Meiji University, Faculty of Law 1881

Meiji University, Faculty of Law 1881 Photo: Meiji University

A number of Western-style brick buildings of the Meiji era have been designated as an important cultural assets and turned into museums, such as the Kyoto National Museum. Other examples may be seen below:

Craft Gallery of the National Museum of Modern Art, built by Hisashi Tamura in 1910.

Craft Gallery of the National Museum of Modern Art, built by Hisashi Tamura in 1910. Designed as “The former Headquarters of the Imperial Guards.” (Photo: Craft Gallery of the National Museum of Modern Art)

Kyoto National Museum designed by Katayama Tokuma

Kyoto National Museum designed by Katayama Tokuma in 1895, and built to house art treasures donated by the Imperial Household Ministry as well as those owned by temples and shrines (Photo: Kyoto National Museum)

Ryounkaku (a.k.a. Asakusa Twelve Storeys), built in 1890, was Japan’s first western-style “skyscraper”, and the first to install an elevator. At 52 meters with 12 storeys, it was the tallest building in Tokyo. The building contained 46 stores selling goods from across the globe.

Ryounkaku (Asakusa Twelve Stories), the first Skyscraper

Ryounkaku (Asakusa Twelve Stories), 1890. (Photo: Heritage of Japan)

Japan had witnessed the improvements in transportation, communications, trade and banking made in the West, where the Industrial Revolution had taken place in the late 1700s beginning in Britain and lasting through till around 1840. Japan wanted to resist the power and hegemony of the West by emulating the West. The Japanese sought Western technology and modernized essentially to level the playing field so they would not be colonized or taken advantage of by the West.   Under the nationalistic slogan “rich country, strong military,” the Japanese government was intent on learning the secrets of the West and Western experts were brought to Japan and Japanese experts were sent abroad to learn everything they could. The government’s top-down modernization policy dictated the goal of “more production through new industry” so that giant government-operated factories for the military, such as the Koishikawa arsenal factory,  were built. The rapid development of industries in Tokyo, besides giving rise to urban problems such as noise and pollution from the smoke emissions from factory chimneys, as well as the formation of labour unions and social unrest with workers protesting their long hours, low wages and sweatshop conditions. Ultimately, that tenacious onslaught and monentum of industrial development both supported as well propelled Japan headlong into two wars: the Sino-Japanese War(1894-1895)  and the Russo-Japanese War(1904-1905).

Koishikawa Arsenal Factory

Koishikawa Arsenal Factory  (Photo credit: Edo-Tokyo Museum; Heritage of Japan)

 Nirayama-Hansharo

In 2015, “Sites of Japan’s Meiji Industrial Revolution: Iron and Steel, Shipbuilding and Coal Mining” mainly located in the southwest of Japan were designated UNESCO World Heritage Sites.

The development of the iron and steel industry, shipbuilding and coal mining underpinned the rapid industrialization of the country from the middle of the 19th century to the early 20th century. To showcase the nation’s achievements, the country’s first National Industrial Exhibition was held on August 21, 1877 (bottom right photo: color print depicting the Opening ceremony to the first National Industrial Exhibition. Credit: National Diet Library). The national event (inspired by the 1873 Vienna International Expo) emphasized its aspect as an industrial promotion opportunity providing a meeting place for Western technologies and their Japanese counterparts, for conducting product survey and industry promotion.

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Colour prints of the first National Industrial Exhibition (Photo credit: National Diet Library)

The approximately 100,000 m2 venue contained the Fine Art Building (top left photo, photo: National Diet Library), the Agricultural Production Building, the Machinery Building, the Horticultural Building, and the Animal Building. An approximately 10 m high American-style windmill (for pumping up groundwater) was constructed at the park entrance. All the exhibits collected from across Japan were categorized roughly into six groups (mining and metallurgy, manufactures, fine art, machinery, agriculture, and horticulture). They were judged based on the criteria of materials, manufacturing methods, quality, adjustment, effectiveness, value and price. Medals, certificates of merit and other honors were bestowed on excellent exhibits

Redbrick warehouses (similar to those that proliferated in Manchester during U.K’s Industrial Revolution), were constructed along with expansion of harbour facilities in the late 19th century by the Yokohama city government.  Planned by a Japanese architect and government official, Tsumaki Yorinaka, the redbrick warehouses were erected in 1911 and 1913 to serve as custom houses. When the 1923 Great Kantō earthquake struck Yokohama, the Akarenga red brick buildings suffered less damage than other buildings due to their reinforced structure with iron implanted between the bricks.

Akarenga redbrick warehouses in Yokohama were completed in the period straddling the Meiji and Taisho periods

Akarenga redbrick warehouses in Yokohama were completed in the period straddling the Meiji and Taisho periods  (Source: Free domain)

Other than buildings, notable Meiji era structures included warehouses(see photo above), lighthouses and aqueducts/bridges, an early example of which is the Suirokaku Aqueduct along the Biwakososui River in Kyoto.

Suirokaku Aqueduct, Biwasosui River, Kyoto, 1890

Suirokaku Aqueduct, Biwasosui River, Kyoto, built in 1890. Tanabe Saburo was only 22 years old when he designed and built the aqueduct.

Eight lighthouses were built according to western specifications during the Meiji era. The lighthouse seen in the photo below is one of eight built during the Meiji period.

Oldest Western-style lighthouse, in Shinagawa, Tokyo Bay waterfront, built by French engineer Francois L. Verny in 1870. The lighthouse is one of eight built during the Meiji period (photo credit: Flicker Creative Commons, Daniel Rubio)

Oldest Western-style lighthouse, in Shinagawa, Tokyo Bay waterfront, built by French engineer Francois L. Verny in 1870. (photo credit: Flicker Creative Commons, Daniel Rubio)


Facial tattoos of the Jomon, and what they may have been for

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Below are a few sample closeup photos of tattooed faces of the Jomon mostly of clay figurines (called dogu) or of pottery relief:

Relief face in a clay pot from Yamanashi prefecture(Minami-Alps city) 3,000BC)

Dogu head from Saga prefecture, 3,300BC

Tattooed face of a dogu from Tokyo (Tama New Town), 3000BC

Dogu from Iwate prefecture, 1300BC

Dogu from Yamanashi prefecture, 3000BC

A sketch of a dogu from Kanaaraizawa site, Ibaragi prefecture

Bearded dogu from Goto site, Tochigi prefecture

tattooed dogu face from Gunma prefecture

Dogu from Yamanashi prefecture (minami-Alps city), 3000BC

Dogu with heavily tattooed eyes and lips from Akita prefecture, Final Jomon

National Treasure dogu figurine from Aomori prefecture, Final Jomon

Dogu from Akita prefecture, Final Jomon

Clay mask from Akita prefecture (Late Jomon Asao site)

Another clay mask showing ornate tattoo markings from Iwate prefecture (Shidanai site), Final Jomon

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Dogu from Chiba Prefecture (Late Jomon period)

A rare figurine with ornate face markings and large headgear, from Ishikawa prefecture, Late Jomon period

Dogu from Ibaragi prefecture, Final Jomon

Dogu from Nagano prefecture, Middle Jomon period

“Mimizuku dogu” from Chiba prefecture, Final Jomon period

“Mimizuku dogu” from Saitama prefecture

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Dogu from Kanagawa prefecture, Late Jomon period

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Dogu from Kanagawa prefecture, Middle Jomon period

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Dogu from the Ikawazu Shellmidden site in Aichi Prefecture (Late Jomon period)

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Clay mask from the Sanganchi Shellmidden site in Fukushima prefecture (Final Jomon period)

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Dogu from Miyagi prefecture (Late Jomon period)

Dogu from Niigata prefecture (Nagaoka city) 3000BC

The most common type of tattoo appears to a simple double line running in a curve or straight line from the eye across the cheek to the side of the face. Other figurines feature crow’s feet lines from the corner of each eye (pictured below).

Dogu from Akita prefecture, 600BC

Eyes and lips may also have completely been surrounded by scarification or needle punctures creating the famous goggle-eyed and thick-lipped look of many figurines.

A closeup of these clay dogu of body parts – lips give an idea of how highly tattooed lips may have been

Eyelids may have been tattooed as well.

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The eyelids appear to have dotted lined tattoos, both upper and lower ones. Clay mask from Aomori prefecture (period unknown)

Why did the Jomon tattoo their faces?

Local fishing folk believe that tattoos are done for protection against predators like shark. This practice may have been handed down from Jomon times.

Some of the tattoos such as the double lines from eye across cheek appear to have been fairly stable features, lasting from around 3,300 BC through the final phase of the Jomon period in the Kanto region. They may have been rites of passage or cultural traits intended to have been cultural/tribe identifiers.

Ainu women were known (until fairly recent times) to have tattooed their lips. According to Ainu tradition, they attributed tattooing to a gift from the “Ancestral Mother” of the Ainu Okikurumi (Turesh) Machi, the younger sister of the creator god, Okikurumi. And (like with the Inuit people)the tattoos were a prerequisite for Ainu women to getting married (source: Lars Krutak). Other Ainus have attributed a healing or protective function to tattoos that are believed to repel evil spirits(source: Lars Krutak). As the Ainu have been established to be genetically partially of Jomon stock, we can postulate that the practice may have been part of the goddess culture of the Jomon people as well (other cultures like InuitEskimo Imooyok (see photo below) and Maoris women also tattooed their lips. Facial tattoos are especially sacred for the Maoris, a moko on the face is the ultimate statement of one’s identity as a Māori, and the head is believed to be the most sacred part of the body).

「tattoos orochi ulchi amur」の画像検索結果

Later periods also showed up more ornamental and varied types of tattoos, where perhaps they became fashion or artistic adornments as well.

Jomon tattoos were unlikely to have had warpaint uses, i.e. they were probably not intended to scare off their enemies. Internecine warfare was unknown until the arrival of the Yayoi immigrants

Sources and References

縄文の力 THE POWER OF JOMON (別冊太陽 日本のこころ Author: 小林 達雄 ISBN9784582922127

土偶・コスモス(Published by MIHO MUSEUM) ISBN9784904702376

列島の考古学 縄文時代 (by 能登) ISBN9784309714424

縄文人の道具 古代史復元 (3) Author: 達雄, 小林) ISBN4061864238

縄文のわざと道具 縄文時代1 (日本のあけぼの)Author: 小林 達雄 ISBN4620603023

 

Tattoo History: The Tattooed Ainu Women of Japan by Minerva

Tattooing among Japan’s Ainu people by Lars Krutak

Ta moko: The significance of Maori tattoos 

Between the Lines:
Tracing the controversial history and recent revival of Inuit facial tattoos
Sep 20, Ashleigh Gaul

‘This is so powerful:’ Kitikmeot women revive traditional Inuit tattoos CBC News,May 3, 2016

The Exact Opposite Idea of the Pirate Cliché

The Exact Opposite Idea of the Pirate Cliché

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A thoughtful piece from the Pirates of Asia blog that lends insights to why the Murakami pirates were such a force to be reckoned, their innovations and how they contributed to the flourishing local island trading economies…

PIRATES OF ASIA

Wokou14n15thc By The original uploader was Yeu Ninje at English Wikipedia – Transferred from en.wikipedia to Commons., CC BY-SA 3.0, Link

The Murakami Pirates were a bunch of ordinary people that turned into a remarkable pirate group, during the late sixteenth century in Japan. These pirates did not do what ordinary pirates did. Instead of assailing other pirates, they were protecting their customers for a living. Eventually they became the greatest pirates in Japan.

The Murakami Pirates had travelled to and settled on some of the islands of the Seto Inland Sea. The Seto Inland Sea is a waterway dotted with some 3,000 islands and islets. The pirates chose that area, so they could have their headquarters. There were three key islands chosen for their base: Noshima, Kurushima, and Innoshima. Seto Inland Sea is right between western Honshu, and northern Shikoku regions.

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It is evident, from viewing the map, that…

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Mori Motonari, the smartest sea lord in Japan

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1000px-Battle_of_Miyajima-2The Island of Miyajima is better promoted and known for its serene scenic beauty, but it is historically significant for one of the most dramatic battle scenes of the Sengoku era, the Battle of Miyajima, which involved drastically outnumbered forces, pirate action and an amphibious castle assault in the crazy cover of a wild storm. Read about it here through the reblogged post by the Pirates of Asia blog…

PIRATES OF ASIA

When I was little I used to play games involving pirates, ships etc but have you, ever wondered how the real pirates looks like? Or do they have the same type of personality like in pirate games? Or are they bad criminals like how people think? Well, I will be talking about a sea lord who controlled a band of pirates to achieve his political ends, and who is a famous hero in Japan.

180px-noshima-murakami-takeyoshiMori Motonari

The painting above shows how he looked. Not pirate-like or angry at all. He was said to be a diplomat and to like poetry. In other words, he was rather gentleman-like and cultured and had manners..

Mori Motonari(毛利 元就) was a feudal lord (1497~1571) in the western Chugoku region during sengoku period of the 16th century. The Mori clan claimed descended from Ōe no Hiromoto (大江広元), an adviser to Minamoto Yoritomo. Motonari is known…

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A map of Japan made during the Italian Renaissance

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In 1587, Italian cartographer Urbano Monte created a world map with a unique perspective centred in the North Pole. This map is from one of the 60 pages of that work. Monte’s knowledge of Japan was apparently sourced from a Japanese delegation he had met in Milan, Italy in 1585. Monte is said to have laid out the islands in a horizontal fashion, according to his unique perspective, which differs from that of the modern globe we are used to. Credit: LIVESCIENCE

A walk on Mt. Mitake brings encounters with wolf and mountain lore

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Musashi-Mitake-jinja

The Musashi-Mitake-jinja Shrine is believed to have been founded in 90 BC by Emperor Sujin, which makes it one of the oldest in Japan. However, the shrine also records that Priest Gyōki built a hall on this site and enshrined the Zaō Gongen statue in it in 736. The mountain, close to Tokyo, has been known as a sacred mountain and an object of worship since ancient times.

Musashi country (Musashi no kuni) painted by Ryusai Yukinobu Source: Great- Edo database

Hinting of the antiquity of the mountains’ ritual practices, the Hinode (Sunrise) Festival is the annual spring festival of Musashi-mitake Jinja Shrine. It is the most important of this shrine’s festivals.

The name of this festival comes from a ceremony held when reaching the peak of the mountain by Buddhist mountain priests, which is thought to be the origin of the 2000-year-old festival, which was supposedly held together with the sunrise on February 8 according to the old lunisolar calendar. These days the festival is held on May 8. (See photo of sunrise on Mt Mitake here.)  Despite the Shinto jinja’s presence being thought of as Shinto, we can see how the syncretism of faiths, of Shinto and Buddhism practices, was at work and was the norm from early times.

The Festival on May 7 includes a present-day solemn Shinto procession starts from the plaza in front of the cable car station on the mountain’s summit and winds through the Mitake village. Mainly focused on a mikoshi (a palanquin-styled portable shrine), this procession also includes Shinto priests, men dressed in white, armored warriors, and children.

Click to view slideshow.

After passing through the village, the procession goes up 300 stone steps to the main shrine at the summit of Mt. Mitake (929 meters high). While the initial procession ends at the location of the sacred palanquin with the receipt of symbols of the deities accompanied by gagaku (traditional Japanese court music) performed at twilight and the faint light of the lamps. At the end, participants are entertained by a kagura (Shinto music-and-dance) performance at the end.

Mountain ascetics, numerous shugenja visited the mountain especially after the Kamakura Period (1185–1333), when mountain religions proliferated. Mountain ascetics secluded themselves in the mountain to perform purification rites, cleansing themselves ritually under the nearby waterfalls (Nanayo-taki and Ayashiro-no-taki).

From the 17th century, commoners began to visit the shrine, and to accommodate all the pilgrim visitors to Mt. Mitake to pray at Mitake-jinja Shrine, the path to the mountain shrine became dotted with Shukubo temples lodgings. Many of those Shukubo lodges remain open today especially to kendo martial arts practitioners and others here to practise mountain austerities and meditation.

As to the origin of the Japanese solar cult religions, author Jacques H. Kamstra – in his 1967 book, “Encounter Or Syncretism: The Initial Growth of Japanese Buddhism” has made strong connections between hi-kami and hinode matsuri practices and the northern Asian continental solar cults possibly via Korean immigrants.

Like most mystical mountains, this one comes with its own set of folklore and legends. But among the folklore, wolves are beloved heroes and not the big bad wolves of modern fairytales. And they are seen in several forms on this mountain, as guardians of the small Inari shrine here, and as venerable guardian Shirookami deity of the Musashi-Mitake jinja, and the sacred messenger that led the legendary imperial ancestor, Yamato Takeru to safety.

Strangely, the baddie in the folklore here is a stag which is honoured elsewhere as a messenger of the gods at Kashima shrine in the east and in Nara’s imperial sheine. According to legend, wolves led Prince Yamato Takeru back to safety after a mountain demon disguised as a stag, led him astray on the slopes of Mount Mitake. Not surprisingly, the prince declared the wolves divine, and that is how they became sacred guardians of the Mitake mountain peak. (One wonders if the stag vs wolf opposition is merely a depiction of totemic animal symbols representing euphemistically an ancient tribalistic rivalry between political factions’ rulers.)

Statue of Hatakeyama Shigetada

The statue of the Kamakura period warlord-Shigetada Hatakeyama on horseback adds an air of samurai splendour to the already colourful shrines. The shrine’s treasure house is also famous for the Akaito Odoshi Yoroi (red-thread-armour) offering (now designated a national treasure) dedicated by Hatakeyama Shigetada in 1191.

Shrine visitors are set to increase in the zodiac Year of the Dog as in 2018. Lots of votive emas of the wolf-dog will be offered up.

Votive ema of wolf

You can view the rest of the photos of my walk on Mt Mitake at Field trip: Following the trail of Mt Mitake’s mountain pilgrims

Finally, if you are planning a trip yourself, you can find information on access to the mountain, by clicking on this page and this page.

ALDH2 and migration patterns into Japan

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Hu Li et al.,2009, Refined Geographic Distribution of the Oriental ALDH2*504Lys (nee 487Lys) Variant

Summary
Mitochondrial aldehyde dehydrogenase (ALDH2) is one of the most important enzymes in human alcohol metabolism. The oriental ALDH2*504Lys variant functions as a dominant negative greatly reducing activity in heterozygotes and abolishing activity in homozygotes. This allele is associated with serious disorders such as alcohol liver disease, late onset Alzheimer disease, colorectal cancer, and esophageal cancer, and is best known for protection against alcoholism. Many hundreds of papers in various languages have been published on this variant, providing allele frequency data for many different populations. To develop a highly refined global geographic distribution of ALDH2*504Lys, we have collected new data on 4,091 individuals from 86 population samples and assembled published data on a total of 80,691 individuals from 366 population samples. The allele is essentially absent in all parts of the world except East Asia. The ALDH2*504Lys allele has its highest frequency in Southeast China, and occurs in most areas of China, Japan, Korea, Mongolia, and Indochina with frequencies gradually declining radially from Southeast China. As the indigenous populations in South China have much lower frequencies than the southern Han migrants from Central China, we conclude that ALDH2*504Lys was carried by Han Chinese as they spread throughout East Asia. Esophageal cancer, with its highest incidence in East Asia, may be associated with ALDH2*504Lys because of a toxic effect of increased acetaldehyde in the tissue where ingested ethanol has its highest concentration. While the distributions of esophageal cancer and ALDH2*504Lys do not precisely correlate, that does not disprove the hypothesis. In general the study of fine scale geographic distributions of ALDH2*504Lys and diseases may help in understanding the multiple relationships among genes, diseases, environments, and cultures.

Central China Origin of ALDH2*504Lys
The frequency decline from Southeast China to West and North China is quite smooth. The allele frequencies decrease to less than 20% in Southwest and Central China, and to less than 10% in Manchuria, Mongolia, Xinjiang, and Tibet within the broader region of East Asia. In Central Asia and Siberia, beyond the pronounced genetic influence of Han Chinese, the ALDH2*504Lys allele is rare. The allele is also detected in some Iranian populations, which may be explained by diffusion along the Silk Road. We conclude that the spread of ALDH2*504Lys to the north and west was concomitant with the expansion of Han Chinese and diffusion of the allele into surrounding populations.
Although the ALDH2*504Lys allele frequency reaches a peak in Southeastern Chinese populations, we cannot draw the conclusion that this allele originated there. The population history shows clearly that Hakka and Minnam Chinese presently in Southeast China are descendants of migrants from Central China (Wen et al., 2004). The indigenous populations in South China, such as Hmong-Mien populations (Hmong and She) from the Yangtze River area, and Daic populations (Kam, Laka, Mulam, and Maonan) from the Pearl River area, exhibit much lower frequency of ALDH2*504Lys. ALDH2*504Lys is almost absent in the aboriginal populations of Hainan and Taiwan, the two largest islands in South China. Therefore, it is unlikely that the Southeast Chinese obtained the ALDH2*504Lys allele from the indigenous populations. Unlike the gradually decreasing frequency to the north and west, the allele frequency drops sharply to the south. The allele exists at low frequency in Peninsular Southeast Asia, and is rare in the Southeast Asian islands. If this allele originated in the Southeast Chinese populations after they arrived in the present region, the quick expansion of the allele to the north and west cannot be explained. Therefore, we conclude that the ALDH2*504Lys allele was most probably carried south by the Han Chinese migrants from Central China, rather than originating in the indigenous populations in the region where it now has the highest frequencies.
Understanding why the present Central China populations exhibit much lower ALDH2*504Lys than the Southeast China populations is crucial in the study of the history of this allele. Both the decrease in Central China and the increase in Southeast China should be accounted for. In the history of China, many Altaic populations moved from the North China to Central China after wars in the 4th, 12th, and 13th centuries which also resulted in the migration of some Chinese populations from Central China to South China. These Altaic populations later merged with the Central Chinese populations after their kingdoms or dynasties ended. The most famous examples are Sienbers (Xianbei, founders of Former Yan Kingdom, Later Yan Kingdom, Western Qin Kingdom, Southern Liang Kingdom and Southern Yan Kingdom of Sixteen Kingdoms Period, and Northern Dynasties), Huns (founders of Han-Zhao Kingdom and Northern Liang Kingdom of Sixteen Kingdoms Period), Khitans (founders of Liao Dynasty), and Jurchens (founders of Jin Dynasty). Those Altaic migrants may have included very few or no individuals carrying the ALDH2*504Lys allele because present Altaic populations have a low frequency of the allele. The merging of these Altaic populations could have decreased the proportion of ALDH2*504Lys in the Central Chinese populations. On the other hand, some as yet unknown protective effects of ALDH2*504Lys against diseases might also have contributed to the increased frequency of this allele in Southern Chinese. Since migrations to South China resulted from wars, the refugees may have been subjected to considerable stress and a selective advantage could have had great impact. We can speculate that the ALDH2*504Lys heterozygotes had an advantage because they tended to drink less alcohol or had some other advantage (Chen et al., 1999). The recent appreciation of other metabolic/pharmacologic roles for ALDH2 (Li et al., 2006; Larson et al., 2007; Chen et al., 2008) suggest that if selective factors are responsible for the high ALDH2*2 frequency in East Asia, their nature may be unrelated to the current association with esophageal cancer or ethanol metabolism. Alternative hypotheses of increased resistance to some disease organisms (Enoch and Goldman, 1990;Yokoyama et al., 2001; Oota et al., 2004; Yokoyama & Omori, 2005; Yang et al., 2007; Li et al., 2008) would also explain a clear advantage to heterozygotes. However, statistically positive selection on ALDH2*504Lys cannot be detected using the extended haplotype test (Sabeti et al., 2007) as very low levels of recombination exist in the genomic region of ALDH2 locus (Oota et al., 2004). Other methods suggest positive selection on ALDH2*504Lys (Long et al., 2006).
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Geographic Association with Esophageal Cancer Incidence
Whatever positive selection may have increased the frequency of ALDH2*504Lys, serious diseases such as esophageal cancer or ischemia could act to decrease the ALDH2*504Lys allele frequency among the populations since studies report that heavy alcohol drinkers who are heterozygotes for ALDH2*504Lys have higher risk for esophageal cancer (Yokoyama & Omori, 2005; Yang et al., 2007; Li et al., 2008). In addition, ALDH2 activation was shown to reduce ischemic damage to the heart, suggesting that patients with reduced ALDH2 activity may suffer increased damage during cardiac ischemic events or coronary bypass surgery (Chen et al., 2008). The typical age of onset for esophageal cancer in the high incidence area can be earlier than 30 (He et al., 2006). We compared the geographic distribution of esophageal cancer incidence with the ALDH2*504Lys allele and carrier frequency distributions. We collected the male esophageal cancer incidence data of 355 populations from the literature, covering most countries in the world (Table S2). Central and Southeast China were examined in detail. Figure 3 illustrates the world distribution of esophageal cancer incidence and the details in East Asia. The extremely high incidences only appear in East Asia and some populations in Central Asia where the frequency of ALDH2*504Lys carriers is also high. However, comparison of Figure 2 and Figure 3 shows that the distributions are far from identical. However, the high cancer incidence areas mostly fall into the high frequency area of the derived allele carriers. The acetaldehyde accumulation resulting from ALDH2*504Lys in those who drink alcohol is certainly not the only risk factor for esophageal cancer. As noted above, ALDH2 also has other metabolic functions that could be independently influencing the distribution of ALDH2 variants (Li et al., 2006; Larson et al., 2007). Some environmental factors such as soil and vegetation characteristics and life styles may also be associated with the esophageal cancer risk (Wu et al., 2007; Fan et al., 2008; Moradi, 2008).

In Central Chinese populations, the heritability of esophageal cancer is estimated at around 49% (Han et al., 1994; Li et al., 1998). East Asian migrants in America also have a much higher esophageal cancer incidence than European Americans and African Americans (Parkin et al., 1997), indicating the pronounced heritability of esophageal cancer. Therefore, the incidence of esophageal cancer is affected by multiple factors that interact with the ALDH2*504Lys allele frequency in a complex way. That complexity could explain the differences between the distributions of esophageal cancer and the ALDH2*504Lys allele carriers in East Asia.
In most areas of South China and Southeast Asia, the incidence of esophageal cancer is much lower than that observed in Central China, indicating that there are fewer environmental risk factors and lower susceptibility of esophageal cancer in South China. However, there is still a high incidence area in Southeast China, which might be associated with the highest allele frequency of ALDH2*504Lys in exactly the same geographic area. In contrast to the high incidence of esophageal cancer in Southeast China being the consequence of the high ALDH2*504Lys frequency, it is possible that the high incidence of esophageal cancer in Central China is working to decrease the ALDH2*504Lys frequency while cultural pressure to consume ethanol increases as the impact of *504Lys decreases. The answer depends on which factors increasing risk are most important in which area and how they interact.
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Conclusion
In conclusion, we hypothesize that the oriental ALDH2*504Lys variant might have originated in the ancient Han Chinese population in Central China and spread to most areas of East Asia with the expansion of Han Chinese and their genetic influences on neighboring populations over the past few thousand years. Some diseases such as esophageal cancer show a complex relationship with the frequency of ALDH2*504Lys. Where the ALDH2*504Lys frequency is high for whatever reason, as in Southeast China, there is a clear increased risk of esophageal cancer in heterozygotes that results in higher esophageal cancer incidences in some subregions. In other areas of China there is also an increased risk of esophageal cancer in heterozygotes (Yang et al., 2007; Wu et al., 2001; Chen, 2005; Yang, 2005; Xiao, 2007) but the lower frequency of ALDH2*504Lys is not sufficient to explain the high incidence of esophageal cancer. More genetic epidemiological investigations in China are required to reveal any possible reciprocal relationship between esophageal cancer and the ALDH2*504Lys allele and identify the other risk factors that appear to be present.

ALDH2, ADH1B, AND ADH1C GENOTYPES IN ASIANS: A LITERATURE REVIEW   Non-pDF version

2004 The evolution and population genetics of the ALDH2 locus: Random genetic drift, selection, and low levels of recombination

Effects of Worldwide Population Subdivision on ALDH2 Linkage Disequilibrium

Full article pdf
Yayoi Source of ALDH2 https://upload.wikimedia.org/wikipedia/commons/8/8b/ALDH2_rs671_genotype_frequency.png

Origins of Japanese lecture by Saitou https://ocw.u-tokyo.ac.jp/lecture_files/gf_14/4/notes/en/04saito_eng.pdf

2017 Takeuchi et al., The fine-scale genetic structure and evolution of the Japanese population

See also Tracing Jomon and Yayoi ancestries in Japan using ALDH2 and JCV virus genotypes

It’s time to embrace northern route dispersal scenarios for the migration of modern humans

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Many voices and new models disproving the Southern Dispersal route model (following the coast of India) have been steadily coming onstream for a while, but it has been difficult to dislodge the consensus model for more than a decade ever since it emerged, and the mountain of media and literature that followed the original model (see Vincent Macaulay; et al. (13 May 2005), “Single, Rapid Coastal Settlement of Asia Revealed by Analysis of Complete Mitochondrial Genomes; Vol. 308. no. 5724”, Science Magazine, 308 (5724), pp. 1034–36, doi:10.1126/science.1109792, PMID 15890885”. These newer models suggest a pincer model including a route that goes either goes north or south of the Himalayas, before arriving at the coasts of South East Asia and northeast Asia.

Below we survey some of the literature representing the dissonance with the Southern dispersal hypothesis, the most recent of which being:

Nicolas Zwyns Et al., The Northern Route for Human dispersal in Central and Northeast Asia: New evidence from the site of Tolbor-16, Mongolia
Scientific Reports, volume 9, Article number: 11759 (2019)

Abstract

The fossil record suggests that at least two major human dispersals occurred across the Eurasian steppe during the Late Pleistocene. Neanderthals and Modern Humans moved eastward into Central Asia, a region intermittently occupied by the enigmatic Denisovans. Genetic data indicates that the Denisovans interbred with Neanderthals near the Altai Mountains (South Siberia) but where and when they met H. sapiens is yet to be determined. Here we present archaeological evidence that document the timing and environmental context of a third long-distance population movement in Central Asia, during a temperate climatic event around 45,000 years ago. The early occurrence of the Initial Upper Palaeolithic, a techno-complex whose sudden appearance coincides with the first occurrence of H. sapiens in the Eurasian steppes, establishes an essential archaeological link between the Siberian Altai and Northwestern China. Such connection between regions provides empirical ground to discuss contacts between local and exogenous populations in Central and Northeast Asia during the Late Pleistocene.

Introduction

Although models for H. sapiens’ early dispersals out of Africa emphasize a southern route to Asia1,2,3,4,5, Neanderthal and Modern Human (MH) fossils in Siberia6,7,8,9 suggest that at least two other dispersals took place across the Eurasian steppe north of the Asian high mountains. Given the size of the area considered, human fossils are few but recent studies have suggested that a major change in the regional archaeological record could be indicative of a large-scale human dispersal event. Known as the Initial Upper Palaeolithic (IUP), it refers to the sudden appearance in contiguous regions of a specific blade technology sometimes associated with bone tools and ornaments10,11,12,13,14,15,16,17. How old these assemblages are, and how long the phenomenon lasts are still controversial questions, and little is known about the timing and environmental context of these population movements. Here we present new data following excavations of an archaeological site located in a low altitude pass, the Ikh-Tolborin-Gol, which connects Siberia with northern Mongolia. Our results document the early occurrence of the Upper Paleolithic in Mongolia and provide a chronological reference for population movement across Northeast Asia during the Late Pleistocene. …

To summarize, the data at hand suggests that H. sapiens are more likely to be the makers of the IUP. The sudden appearance of this technology in Mongolia takes place in GI12, a temperate phase contemporaneous with the expansion of H. sapiens, in Western and Northern Siberia. Other scenarios cannot be rejected but they are less parsimonious, and they simply lack convincing alternatives regarding the archaeology of H. sapiens populations. Either way, the population movements illustrated by the dispersal of the IUP occur in a general context of climatic instability associated with MIS3 in Mongolia, and more generally in Central and Northeast Asia66. Such variations in the higher latitude of the Eurasian steppe imply different challenges (eg. continental climate, aridity) than those met along the southern route into the Indian subcontinent67. An inland route to Asia borne out by the archaeological and fossil record is, however, not exclusive of other dispersal models (e.g. Southern or ‘coastal’ Route) while it remains consistent with the available genetic data. The genome of an aboriginal Australian shows that deeply rooted populations have a shared ancestry coming from at least two early H. sapiens dispersals out of Africa into Asia68. Meanwhile, extant populations from continental East-Asia69 and in Melanesia70,71show evidence of gene flow from Denisovan individuals from at least two gene flow events65. The timing might still be uncertain, but the dates presented here for the dispersal of IUP are consistent with the age estimates for an early H. sapiens – Denisovan encounter, distinct from the other gene flow events that would take place much later in Southeast Asia (or Melanesia)65. After subsequent mutations, the genes passed along from Denisovans around 45 ka may have been decisive for the survival of our species in extreme environments (Tibetan populations’ adaptation to hypoxia of high altitude)72,73. Finally, it is notable that East-Asian and Melanesian populations do also show evidence of gene flow coming from Neanderthal individuals65. Granted that the Pleistocene population dynamics were probably complex, a model integrating dispersals of several taxa, such as Neanderthal and H. sapiens across the continent is a parsimonious complement to a strict coastal migration scenario74, 42.

Conclusions

The results obtained at T16 indicate that the IUP techno-complex occurs in Western front of East Asia as early as 45 ka. Contemporaneous with skeletal evidence of H. sapiens, the IUP in Tolbor establishes an essential archaeological link between Siberia and Northern China. Together with evidence from these adjacent regions, the material found in AH6 suggest that a widespread behavioral shift took place along the Eurasian steppe belt during a period of climatic instability.

Corollary scenarios that extend the northern route scenario theories are now more plausible ones — see various authors‘ writings on the northeast Asian/East Eurasian origins of Tibetan/Nepali/Sherpa populations, and  George Van Driem(The Eastern Himalayan corridor in prehistory” whose model (see maps immediately above)for a Trans-Himalayan Ice Age Refugia”route, and Larruga et al., of the following 2017 paper:

Carriers of Mitochondrial DNA macrohaplogroup R colonized Eurasia and Australasia from a Southeast Asia core area:

Coeval independently dispersals around 50 kya of the West Asia haplogroup U and the Wallacea haplogroup P, points to a halfway core area in southeast Asia as the most probable centre of expansion of macrohaplogroup R, what fits in the phylogeographic pattern of its ancestor, macrohaplogroup N, for which a northern route and a southeast Asian origin has been already proposed.

Viral markers model migration showing hitherto unknown northern routes

Proposed routes of migration carrying JCV genotypes to the New World. It is assumed that the initial dispersal of JCV from Africa coincides with ‘Out of Africa 2’, but JCV phylogeny does not provide any independent estimate of the dates. Dispersal of some genotypes with the earlier ‘Out of Africa 1’ migration is still a theoretical possibility. Later migrations carried distinctive genotypes to island southeast Asia and Oceania. The Americas, however, have the northeast Asian genotype, type 2A. Post-Columbian migrations from Europe (types 1 and 4) and forced migrations from Africa (types 3 and 6) to North and South America complete the picture. It should be noted, however, that no studies of JCV in African populations in Brazil have yet been reported.

– Chaubey et al., had in 2012 proposed that AA speakers in India today are derived from dispersals from southeast Asia, followed by extensive sex-specific admixture with local Indian populations, disproving the timing of dispersal from a westerly and Indian coastal source. Source: Population genetic structure in India Austroasiatic speakers: The role of landscape barriers and sex-specific admixture

— This paper(excerpted below) supersedes an earlier paper showing a counterclockwise dispersal of the N Mtdna macrohaplogroup from southern East Asia, and is more consonant with the large density of N across Eurasia: Carriers of Mitochondrial DNA Macrohaplogroup N Lineages Reached Australia around 50,000 Years Ago following a Northern Asian Route

Fig 1. Geographic dispersal routes of (A) AMH out of Africa migration, and (B) secondary worldwide human expansions, deduced from the age and geographic localization of L3 and N(xR) mtDNA haplogroups including Lineages O and S from Australia. Climatic marine isotope stages (MIS) and most probable places of genetic admixture with Neanderthals and Denisovans are depicted.

Basic N haplogroup trees (excepting N1kt, N2, N3, X and A) showing coalescence ages and sample geographic origin.

Practically all humans out-of-Africa belong to mtDNA macrohaplogroups N or M, both sister branches of L3 African clade. N shows a global Eurasian distribution but most of its lineages everywhere are members of the R subclade. Only in Aboriginal Australians N(xR) lineages reach frequencies over 50% [5,79], and in some regions of East and Central Asia, haplogroups N9 and A can, respectively, exceed 10% [30,39,58,68,80]. In the rest of its geographic range, the presence of N(xR) lineages is residual and represent small younger expansions driven by the later spread of human groups, mainly harboring R derivatives in Western Asia and R and M derivatives in South and East Asia. …

N5 could not be an Indian autochthonous clade based on phylogenetic reconstruction, as it shares transition 1719 with its sister clade N1 that is a haplogroup of undoubted West Asian origin [34], and it has been also found in the Caucasus, Pakistan, Iran and Nepal [26,6971]. Third, other N lineages detected in India as I, W, X2, or N9a, Y2 and A4, are derived branches of the basal clades N1, N2, X, or N9 and A, of western and eastern Asian origins respectively. Thus, the presence of N lineages in India is better explained as the product of late migration from northwestern and northeastern areas. Even though haplogroup N5 is accepted as an autochthonous Indian lineage, its coalescence age (35.7 ± 8.2) is significantly younger (p < 0.0055) than that of the Australian S lineage (46.8 ± 5.5). This scenario strongly contrasts with the huge presence of autochthonous M [40,72] and R [31,59,60,73] lineages with deep coalescence ages in India. It could be alleged that primary autochthonous N lineages existed in India but became extinct due to genetic drift, but this hypothesis is in contradiction with the fast population growth detected in prehistoric southern Asia [74]. In summary, it seems that the first colonizers of Australia, carrying mtDNA haplogroup N(xR) lineages, could use a route not involving India as a stage. …

Our phylogenetic and phylogeographic analysis of macrohaplogroup N in Eurasia supports the existence of an additional northern route out of Africa, not involving the Arabian Peninsula or the Indian subcontinent as previously envisaged [17]. This long journey ended in Australia when it was still a part of the Sahul, most probably at the last glacial stage MIS-4 (Fig 1A). On the top of the common L3* trunk, macrohaplogroup N accumulated a stem of five mutations without any known bifurcation. From this fact, it can be deduced that, after the out-of-Africa, the bearers of this lineage seem to have had demographic difficulties and remained as a stagnate population for a long time. So, the first stages of the proposed haplogroup N northern route would be speculative and have to find indirect support on other genetic, archaeological and anthropologic evidences. … as this study demonstrates, an additional Levant northern route is more congruent with available multidisciplinary data. In addition, combined genetic, archaeological and bioclimatic evidence suggest that, although the early anatomically modern human was born in Africa, the nursery of the modern humans that colonized Eurasia, Oceania and the New World might be first at the south Siberia northwest China core and later in Southeast Asia.

— Candidates for Hoa binhian (early dispersals in Asia) are found in Yunnan, SW China (thought to be bearers of Mtdna R9) which may indicate that the previously-supposed southern early people populating SEA may have had a more northerly origin than thought.

— Excerpts from the Abstract and body of the paper by Sheila Misora et al., Continuity of the microblade technology in the Indian Subcontinent since 45ka: Implications for the dispersal of modern humans

… We extend the continuity of microblade technology in the Indian Subcontinent to 45 ka, on the basis of optical dating of microblade assemblages from the site of Mehtakheri, (22° 13′ 44″ N Lat 76° 01′ 36″ E Long) in Madhya Pradesh, India. Microblade technology in the Indian Subcontinent is continuously present from its first appearance until the Iron Age (~3 ka), making its association with modern humans undisputed. It has been suggested that microblade technology in the Indian Subcontinent was developed locally by modern humans after 35 ka. The dates reported here from Mehtakheri show this inference to be untenable and suggest alternatively that this technology arrived in the Indian Subcontinent with the earliest modern humans. It also shows that modern humans in Indian Subcontinent and SE Asia were associated with differing technologies and this calls into question the “southern dispersal” route of modern humans from Africa through India to SE Asia and then to Australia. ….

The association of microblade technology with modern humans in the Indian Subcontinent is undisputed due to its continuity up to around 3 ka. …Blade technology which has a sporadic but early appearance in the Middle East and Africa [9,10,11,12] is absent from the Indian Acheulian and Middle Palaeolithic. Projectile technology which was an important development in Africa and Europe [13,14] during the post Acheulian period is also virtually absent from the Indian Middle Palaeolithic [15,16]. This suggests that microblade technology is not indigenous to the Indian Subcontinent. Mehtakheri is currently the oldest dated microblade site in the Indian Subcontinent and extends the origin of this technology in the Indian Subcontinent to 44 ± 2 ka based on the weighted average of four dates and ~ 48 ka if the oldest of the dates is accepted as the most accurate as argued below. While microblade technology is associated with modern humans in the Indian Subcontinent, this is not so elsewhere, at least until a much later time period. Modern humans in the Middle East are associated with Middle Palaeolithic technology [17], in Sub-Saharan Africa with the Middle Stone Age [18] and in Southeast Asia [19] and Southern China with core and flake industries [20]. Each of these categories themselves encompasses significant variation. …

Based on our older ages for microblade technology in the Indian Subcontinent, and the arguments presented above, we present the following model for modern human dispersal (as illustrated in Figure 2):

  • 1. During MIS 6 widely dispersed hominin populations with a common ancestor in Homoerectus had differentiated into distinct populations with modern humans and possibly other archaics in Africa, Neanderthals in Europe, Denisovans in temperate Eastern Eurasia and archaic Indians in the Indian Subcontinent and Sundaland (Figure 2.1).

  • 2. During the Interglacial climate of MIS 5, modern humans expanded into Eurasia at the expense of both Neanderthals and Denisovans and reached SE Asia. Due to competition with Indian archaics, modern humans were unable to disperse into the Indian Subcontinent and modern humans reached SE Asia at this time via a northerly route through the Middle East, Central and Eastern Eurasia and Southern China. During this time admixture with both Neanderthals and Denisovans is likely to have occurred in the modern humans reaching SE Asia (Figure 2.2) … Sharp cultural boundaries between the Indian Subcontinent and adjacent areas are not in conformity with a rapid dispersal through India to SE Asia from Africa. Archaeological sites in Arabia and Africa dating to MIS 5 show some distinctive technological features, such as Nubian cores and bifacial projectiles which are absent from any assemblage in the Indian Subcontinent, making it unlikely that modern humans dispersed into the Indian Subcontinent at this time. On the other hand microblade technology or blade technology is attested to at archaeological sites dating to around 60 ka in Africa which have a closer resemblance to the Indian microblade technology. Core and flake assemblages are associated with modern humans in SE Asia and may date back to MIS 5 times. We suggest that the later entry of modern humans into the Indian Subcontinent compared to adjacent regions is because Indian Archaics could easily compete with modern humans during climate conditions favorable to both. …

– A number of research papers in recent years have been coming onstream, concluding that the findings of variations of Altai Denisovan, Neanderthal genetic introgressions into Asian genes suggest complex migration scenarios that do not corroborate the Southern dispersal route hypothesis(see Tracing the peopling of the world through genomics). Since Neanderthal and Denisovan fossils are located in northerly or central zones, it goes that encounters had to have taken place further north than along the southern Indian Ocean coastal route.  Along with the emergence of a great many ancient DNA with some of the oldest dates seen in northern parts of Eurasia, such as Ust’ Ishim or Mal’ta beret fossils, a model that includes a northern route now seems a lot more plausible. Incidentally, these new scenarios are closer to the findings and proposed migration model of virus markers — compare the map immediately below…

Pavesi found:

Type 2 includes several variants, with subtype 2A mainly in the Japanese population and native Americans (excluding Inuits), 2B in Eurasians, 2D in Indians, and 2E in Australians and western Pacific populations (Fernandez-Cobo et al., 2002; Yanagihara et al., 2002; Zheng et al., 2003; Miranda et al., 2004; Takasaka et al., 2004). Subtype 7A was found to be characteristic of southern China and South-East Asia (Saruwatari et al., 2002), while subtype 7B of northern China, Mongolia and Japan (Sugimoto et al., 2002b; Zheng et al., 2004a). A third subtype (7C), spread throughout northern and southern China, has recently been character- ized by Cui et al. (2004). Finally, type 8 was found in Papua New Guinea and the Pacific Islands (Jobes et al., 2001; Yanagihara et al., 2002).
The ubiquitous distribution of JCV, combined with a transmission mechanism largely within families or popula- tions (Kunitake et al., 1995; Kato et al., 1997; Suzuki et al., 2002; Zheng et al., 2004b), make it an attractive candidate for reconstructing human migrations dating to prehistoric times. The close relationship of JCV found in native Americans with that in North-East Asia is consistent with the migration of Amerindian ancestors from Asia across the Bering land bridge (Agostini et al., 1997a). Doubts regarding the reliability of JCV as a marker of human evolution (Wooding, 2001) have recently been dispelled by a whole-genome phylogenetic analysis focused on the distinction between slow- and fast-evolving sites (Pavesi, 2003). By this approach, it was proposed that the associa- tion of JCV with humans originated in Africa, since type 6 was found to be the putative ancestral genotype. It was also demonstrated how type 6 gave rise to two independent evolutionary lineages: one including types 1 and 4, the other including types 2, 3, 7 and 8 (Pavesi, 2003).

Source: Pavesi, 2005 Utility of polyomavirus in tracing human migrations dating to prehistoric times The Journal of general virology 2005 DOI:10.1099/vir.0.80650-

…See also the pincer models or central Asian route viz. 2012 Alvie’s et al., model below:

Alves I, Šrámková Hanulová A, Foll M, Excoffier L (2012) Genomic Data Reveal a Complex Making of Humans. PLoS Genet 8(7): e1002837. https://doi.org/10.1371/journal.pgen.1002837

See excerpts from the abstract and relevant portions of the paper below

We underline the need to properly model differential admixture in various populations to correctly reconstruct past demography. We also stress the importance of taking into account the spatial dimension of human evolution, which proceeded by a series of range expansions that could have promoted both the introgression of archaic genes and background selection.

Interbreeding between Modern and Archaic Humans

In line with previous studies [10][12]which suggested that some aspects of human genomic diversity were incompatible with a complete replacement of archaic hominins, evidence for admixture between humans and Neanderthals emerged from the first analysis of a complete Neanderthal genome [13]. Indeed, the presence of a significant excess of Neanderthal-derived alleles in Eurasian populations as compared to Africans has been interpreted as resulting from an admixture episode between the ancestors of Eurasians and Neanderthals somewhere in the Middle East [13](Figure 1A). Even though the existence of a very ancient population subdivision in Africa from which both Neanderthals and Eurasians would have emerged could lead to similar patterns [14], the maintenance of such a subdivision over tens of thousands of generations seems unlikely. The sequencing of another archaic hominin from the Denisova cave in the Altaï mountains in Siberia has further revealed that Papua New Guineans showed signs of introgression from this archaic human [15]. Further studies of 33 populations from Southeast Asia and Oceania [16]showed that Denisovan admixture was actually present in other Oceanians, Melanesians, Polynesians, and east Indonesians but was virtually absent in mainland east Asians (but see [17]for evidence of possible Denisovan introgression on the Asian continent). Overall, these genomic analyses of admixture suggest that 1%–3% of the genome of all Eurasians and native Amerindians is of Neanderthal origin [15], and that Papua New Guineans and Australians have another 3.5% of their genome of Denisovan origin [16]. The out-of-Africa model of human evolution, which posited a complete replacement of archaic by modern humans in Eurasia, thus needs to be modified to include a limited assimilation of archaic genes, but the fact that most of the genetic variation observed in extant non-African populations comes from Africa remains true.

…The fact that Denisovan admixture had been first evidenced in Papua New Guineans suggested that admixture had occurred as a single pulse in Southeast Asia, after the separation of the ancestors of Oceanians and other Asians [15], [16](Figure 1A). The analysis of an Australian genome has confirmed the presence of Denisovan admixture in Australians [24]and suggested that admixture occurred during a first early wave of colonization towards Oceania, either in Southeast Asia or earlier in Eurasia (Figure 1B). A reanalysis of a large human SNP database and its comparison with Denisovan-derived alleles has suggested the presence of Denisovan admixture in East Asians, albeit at lower levels than in Oceanians [17], which could have occurred at a different place than for Oceanians, somewhere in East Asia (Figure 1C). Contrastingly, Currat and Excoffier [25]introduced a spatially explicit model of interbreeding between Neanderthals and Eurasians that could occur over the whole Neanderthal range (Figure 1D). They obtained similarly low levels (1%–3%) of Neanderthal introgression in both Europe and China if interspecific exchanges were locally extremely limited (only 200–400 interbreeding events over the >6,000 years of co-existence between the two species). An extension of this scenario to Denisovan admixture would imply that modern humans could have hybridized along all migration routes overlapping with the range(s) of archaic humans (Figure 1D). The fact that the largest levels of Denisovan introgression are found in Oceanians raises the question of a potential discontinuity in the Denisovan range (Figure 1A, 1B) or of a genetic differentiation of archaic hominins living in different ecosystems (Figure 1D). Alternatively, modern humans could have admixed with other hominins [26], and/or inferred hominin introgression could result from the sharing of some derived sites between Neanderthals, Denisovans, and unidentified archaic hominins. A scenario involving an unsampled Eurasian archaic hominin has received support from a recent study [27]showing the presence of a highly divergent (>3 Mya) haplotype of the innate immune gene OAS1. This deep lineage is found at high frequencies in Oceania (and at lower frequencies up to Pakistan). This DNA segment is more closely related (0.6 Mya divergence) to the Denisova sequence than to the Neanderthal sequence, which is itself closer to the human reference sequence. It has been speculated [27]that this fragment had introgressed from a more archaic hominin than Denisovans, who could have been themselves introgressed earlier….

Our understanding of the exact sequence and location of admixture events would highly benefit from a more precise knowledge of the nature and the distribution of Neanderthal segments in our genome. Unfortunately, current estimations of introgression levels are based on a statistic measuring a genome-wide difference in the proportion of archaic-derived alleles between two human populations [13], [14], so that the genomic distribution of introgressed segments is still unknown. However, in addition to the OAS1segment mentioned above [27], several authors have recently argued they had identified candidate regions harboring archaic haplotypes [13], [28], [29]. These regions usually show highly divergent haplotypes with very little evidence for recombination [30]. A dozen genomic regions where Eurasians have haplotypes much more divergent than Africans and a high proportion of derived Neanderthal alleles have been proposed as candidates for Neanderthal introgression [13]. More recently, an X-linked haplotype (B006) in an intron of the dystrophin (dys44) gene, almost absent from Africa but with 9% average frequency outside Africa, has been proposed to be of Neanderthal origin [29]. It is close to the ancestral X haplotype, shares 2/3 of derived alleles with Neanderthals, and has little associated diversity, suggesting a recent origin in humans. Another study has also suggested that several immune-related HLA class I alleles in humans could be of Denisovan origin and that they helped Eurasian populations build their immunity [28]. Whereas the hypothesis of an adaptive introgression is highly seductive, its support is relatively thin. “Denisovan” HLA class I alleles are currently not confined to Oceania but are found widespread in Asia. Moreover, the strongest argued case of Denisovan allelic ancestry (HLA-B*73) is actually not found at all in the Denisovan genome and is presently distributed in western Asia, well in the former Neanderthal range. One should therefore be extremely cautious not to assume that each very divergent haplotype found in humans is necessarily of archaic origin, as cases of incomplete lineage sorting are not rare between higher primates [31], especially in the HLA system where trans-specific polymorphism is facilitated by balancing selection [32]. However, if some introgressed genes were really advantageous, they should have spread and fixed in the human population, but as discussed below there is no widespread signature of strong selective sweeps in Eurasia.

…it is likely that differential admixture should affect population genetic affinities under more complex models of population differentiation. The proper interpretation of human genetic affinities should thus probably be re-evaluated in the light of these results. In particular, the divergence between Africans and Oceanians (showing up to 5% archaic admixture [16]) could be more recent than previously reported (62–75 Kya [24]). It remains unclear whether the method used by Rasmussen et al. [24]to date this divergence is also sensitive to differential introgression, but, if that was the case, the colonization wave to Oceania thought to well predate that towards East Asia [24]could have occurred at roughly the same time once differential admixture had been taken into account.

The paper’s author Clarkson suggests that those early dates of a Middle-Stone-Age-like people are consonant with the previous hypotheses of an early Eurasian split with Papuans (i.e. Abor. Australians’ ancestor) as well as of the incorporation of Neanderthal genes into their lineages. See Chris Clarkson, Richard G. Roberts, Zenobia Jacobs, Ben Manwick, Richard Fullagar, Lee J. Arnold & Qian Hua(2018): Reply to Clarkson et al., (2017), ‘Human occupation of Northern Australia by 65,000 years ago’, Australian Archaeology, DOI:10.1080/03122417.2018.1402884

On the question of whether there was an early arrival in the Japanese archipelago from the north, the controversy is a long-standing one. On the one hand, archaeology has shown the arrival of a northern set of lithics – microblades accompanied by burins (which has only a northern Eurasian transmission route) and virus markers and archaic genes suggest a northern migration route, on the other hand, these thin signals of a northern route taken during glacial times, the evidence of such perilous journeys likely buried in the permafrost, and the voices supporting it are often drowned out by the plethora of denser data on later migration entries from the south. This 2017 paper elucidates the uphill task of proving the early northern route that scientists face:

On the Pleistocene Population History in the Japanese Archipelago | Current Anthropology: Vol 58, No S17, Nakagawa, 2017
This paper provides a current understanding of human population history in the Pleistocene Japanese Archipelago, particularly with respect to the routes and timing of hunter-gatherer migrations, by incorporating multiple lines of evidence from the records of archaeology, human paleontology, and genetic studies. The human fossil remains are concentrated on the Ryukyu Islands in southwestern Japan, suggesting that there may have been a northward migration via the Ryukyu Islands. In contrast, studies of ancient mitochondrial DNA demonstrate genetic continuity among Holocene hunter-gatherer populations in the Paleo-Sakhalin-Hokkaido-Kurile Peninsula, whereas the Pleistocene genetic history is little explored. Although it is largely supported, the assumed population continuity from the Pleistocene to the Holocene inside the Japanese Archipelago is also challenged by an examination of the Paleolithic record and a comparison of the short- and long-term chronologies of the Japanese Paleolithic, implying that the Japanese Paleolithic record was created by hunter-gatherer population migrations from the north and south with substantial time lag and endemic technological invention and transformation during the Late Pleistocene.

“How Do the DNA Studies Tell Us about the Routes of Human Entry into Japan?

The Holocene human fossil record supports an admixture model in which the Paleolithic population originated from both southeastern and northeastern Asia (e.g., Hanihara 1991). The mitochondrial DNA (mtDNA) analyses of modern Japanese revealed that non-African superhaplogroups M and N originally derived from modern H. sapiens dispersing out of Africa (Forster 2004) that eventually came to be the Japanese indigenous populations of Ainu and Ryukuan (e.g., Tanaka et al. 2004; cf. Maca-Meyer et al. 2001). Because Ainu and Ryukuan are descendants of the original Jomon populations (Hanihara 1991; Horai et al. 1996; Omoto and Saitou 1997) and M and N superhaplogroups represent southern and northern routes of human migrations, respectively (Tanaka et al. 2004), the Holocene Jomon population was founded by both northward and southward gene flows.

Studies of ancient mtDNA from the Jomon skeletal remains of Hokkaido show genetic relations between Jomon and Ainu, because both populations retain high frequencies of the haplogroup N9b (Adachi et al. 2011), whereas N9b is scarce among East Asian populations other than Japanese (Tanaka et al. 2004:1842) and is likely skewed to northern regions in Japan (Shinoda 2007). Because the coalescent time of N9b is estimated to be approximately 22,000 year ago (Adachi et al. 2011:355), populations that have this haplogroup emerged around the LGM. In Hokkaido, Epi-Jomon human remains in Hokkaido also have N9b (Adachi et al. 2011), which suggests some degree of gene flow during the LGM to the late Holocene in Hokkaido (22,000–2000 years ago).

As discussed above, both genetic studies based on ancient Jomon mtDNA and those based on modern mtDNA more or less support the “dual-structure model” (Hanihara 1991). This also suggests a complex population history even during the Holocene. Nevertheless, what do these genetic implications tell us about Pleistocene population migrations into Japan? In other words, what does the genetic affinity of the Jomon peoples tell us about Paleolithic population dynamics? In general, because the descendants of Jomon and Yayoi both contributed to the formation of the current Japanese population, Paleolithic foragers should be regarded as the founding population of the Jomon (Hanihara 1991). However, the extent to which Pleistocene Paleolithic populations contributed to modern Japanese is largely unidentified, mainly because there are few genetic and human fossil records, with the exception of some good fossil specimens, notably Minatogawa Man (Baba, Narasaki, and Ohyama 1998; Suzuki 1982). The remaining question is how we understand the complexity in Japanese Paleolithic population history. A question that will not be addressed here is whether there is clear evidence that the Jomon were the direct descendents of the Japanese Paleolithic foragers and whether both hunter-gatherer populations were genetically continuous for the past 30,000 years in Japan.

What Does Archaeology Tell Us about Human Entry into Japan?

The Paleolithic archaeological record provides a basic picture of Pleistocene human population history in Japan. Although the number of Paleolithic sites during the 1960s was only slightly more than 300 (Ohyi 1968:52), the number of registered sites is now greater than 15,000 (Japan Palaeolithic Research Association 2010). Some clarification is necessary, however, regarding this latter number. The “sites” in the recent database include assemblages and collections of artifacts recorded in various contexts, ranging from extensively excavated sites to a few specimens collected on the surface. Because a single cultural level in a deeply excavated multilayered Paleolithic open-air site is counted as a single site, a single location was sometimes counted multiple times, and site size and artifact density from a single site are not standardized among the recorded sites. Although some bias is present in the record, the database is still useful to explore to understand general macro- and microregional patterns of human occupation across the entire Japanese Archipelago.

Considering the regional geographic features and Paleolithic culture history, I divided the 47 prefectures into 7 broader regions (fig. 2). From north to south, they are labeled as north (N), northeast (NE), southeast (SE), central (C), southwest (SW), south (S), and far south (FS). N, S, and FS are isolated islands corresponding to Hokkaido, Kyushu, and Ryukyu islands. NE, SE, C, and SW are the divisions of Honshu Island, the main island in the archipelago along with adjacent Shikoku. Divisions of NE, SE, C, and SW are based on the presence of mountain chains, plains, and the Pleistocene paleogeography. For example, C is the region characterized by high-altitude mountains and plains mostly above 600–1,000 m asl. SW is the region in the middle of the Pleistocene Paleo-Honshu Island. Using the site location data recorded in the database, the number of archaeological sites is counted according to the microregions (table 2). The microregions are then sorted by site density using the areal extent data announced by the Geospatial Information Authority of Japan (2015). SE is the microregion with the highest density, followed by S, C, SW, NE, N, and FS. The highest density in SE is probably explained by sampling bias, due to the high population density in the Tokyo area. It is also because the deeply excavated sites yielded multiple levels of human occupation on the Musashino and Sagamino Uplands in the southern part of SE (e.g., Yajima and Suzuki 1976; Yamaoka 2010). Except for the microregions with the highest density (SE) and lowest density (FS), the site density shows a south to north inclination. High site density in SE, followed by a gradual increase from C to SW, NE, and N, is observed. The sites are all attributed to the Pleistocene, whereas the chronological affiliations of these sites vary depending on the region, especially between N (the southern part of Paleo-SHK) and the rest of the microregions (i.e., Paleo-Honshu and Ryukyuan islands). The Paleolithic in the Paleo-Honshu record started at the beginning of the Upper Paleolithic, around 40,000–37,000 years ago, and ended around 11,500 years ago (Yamaoka 2010; Yoshikawa 2014), whereas the beginning of the Paleolithic record in Hokkaido is not earlier than 30,000 years ago (Izuho et al. 2012; Naoe and Kudo 2014). Thus, the time depth of the Paleo-Honshu Paleolithic record is approximately 27,000 years, as opposed to 18,500 years for Paleo-SHK, because the reliable dates obtained from the hearths in the Agonki-5 site in Sakhalin are 23,500 years ago (Kuzmin et al. 2004; Vasilevski 2003). Because of the difference in time depths, the south to north inclination of site density implies that the earlier Paleolithic sites are more abundant in southern Japan than in northern Japan. High site density in the S microregion (Kyushu) next to the SE of the southern Kanto region in Honshu suggests that waves of the earlier hunter-gatherers would have migrated into Kyushu and spread to the north along Paleo-Honshu Island. Conversely, the likelihood of earlier human population migrations in the early Upper Paleolithic (EUP) from eastern Siberia via Paleo-SHK is not supported. On the one hand, site density patterns alone do not answer the question of timing and size of northerly migratory populations from Paleo-Honshu to Paleo-SHK. The lowest density of the FS microregion of the Ryukyu Islands suggests that human arrivals into the Ryukyu Islands were relatively low and that occupations were not necessarily continuous, unlike the situation in the microregions in Paleo-Honshu. Relatively high site density in C (the central region in Paleo-Honshu) suggests that humans occupied high-elevation regions during the Upper Paleolithic. Good examples are represented by the open-air sites located on the Nobeyama Plateau, where Paleolithic hunter-gatherers could have followed seasonal movements between the central highlands and southern Kanto regions in SE (e.g., Tsutsumi 2011), similar to pastoral transhumance (e.g., Chang and Tourtellotte 1993), and where groups of hunter-gatherers seasonally aggregating to kill large herbivores around lakes would have sometimes succeeded (e.g., Norton et al. 2010b). Given the population entry routes (fig. 1), the observed south to north inclination of site density in the Paleo-Honshu suggests that the majority of Paleolithic migratory groups were from the Korean Peninsula and southern China. If so, routes 1 and 5 are the best-supported routes for early hunter-gatherers’ dispersals into the Japanese Archipelago.

figureFigure 2. Microregions in the Japanese Archipelago. Bold lines represent the boundaries of microregions. Dotted lines within the islands define the current 47 administrative prefectures. Locations with numbers show the human paleontological and/or pre–Upper Paleolithic archaeological sites mentioned in the text. The sites are Minatogawa (1), Yamashita-cho (2), Pinza-abu (3), Shiraho-Saonetabaru (4), Shimojibaru (5), Kanedori (6), Takesa-Nakahara (7), Hamakita (8), Sunabara (9), Iriguchi (10), Sozudai (11), and Ōno (12). C = central; FS = far south; N = north; NE = northeast; S = south; SE = southeast; SW = southwest.

figureTable 2. Counts, areal extent, and density of Late Pleistocene sites in Japan

Paleolithic Chronologies in Japan: Short- versus Long-Term Chronologies

In the  the study of population history, an establishment of cultural chronology is one of the major debated areas of research among the other topics in Paleolithic studies in Japan. Below, I give an overview of the long-term and short-term chronologies and discuss how both chronological models are relevant to global models of human population migrations and dispersals in Eurasia.

Clear evidence of the Japanese Paleolithic appears beginning around 40 ka, and blade technology was incorporated since the earliest lithic assemblages appeared in the southern Kanto region in the SE microregion (Yamaoka 2010; but see Nakamura 2012). The gradual but consistent increase in the number of blade tools (e.g., endscrapers, burins, and perforators) and various blade-production technologies, including prismatic blade technology, which certainly spread across Japan during the Upper Paleolithic, suggests that the technology of the Japanese Upper Paleolithic is not dissimilar to that of the Upper Paleolithic in Europe. On the contrary, unique stone tools characterized in the Japanese EUP are principally represented by three classes of stone tools (fig. 3): trapezoids defined as abruptly and/or minimally retouched small flakes (Sato 1988), backed blades (Ono 1988) characterized by abrupt retouches and truncations on elongated flakes and/or blades traditionally described as knife-shaped tools (Serizawa 1960; Sugihara 1965; Tozawa 1990), and edge-ground axes (Tsutsumi 2012). The combination of trapezoids, knife-shaped tools, and edge-ground axes in EUP is unique to the Japanese Paleolithic industry, and they have not been recovered together in neighboring regions, such as South Korea (K. Bae 2010; Lee, Bae, and Lee 2016), which suggests that they were newly innovated in Japan at the beginning of the Upper Paleolithic; however, edge-ground stone axes attributed to MIS 3 have recently been identified at the Galsanri and Yonghodong sites in Korea (Lee, Bae, and Lee 2016). Indeed, knife-shaped tools long persisted as the formal stone tool class in the Japanese lithic industries, and the “knife-shaped tool culture” is the technocomplex that is extensively distributed from Kyushu to southern Hokkaido (e.g., Ambiru 1986; Morisaki 2012; Naganuma 2010; Ono 1988; Yoshikawa 2010). In the Korean Peninsula, the Upper Paleolithic industry has tanged points (Seumbe Chireugae) from its initial stage with the emergence of blade technology (C. Bae 2017, in this issue; K. Bae 2010; Lee 2015, 2016; Seong 2008, 2009; Seong and Bae 2016). Tanged points also appeared in Japan in the late Upper Paleolithic, around 30,000 years ago, mainly in the Kyushu region; however, they occur rather briefly, perhaps in response to small-scale human migrations from Korea or cultural transmission after the collapse of the regional environment in Kyushu, caused by the large explosive event of the Aira Volcano, which occurred some 30,000 years ago (Matsufuji 1987; Morisaki 2015). The traits shared between the retouch technologies used in the Japanese knife-shaped tools and the Korean tanged points make archaeologists hypothesize that an immediate technological transmission of tanged points from Korea to Japan at the beginning of the Upper Paleolithic stimulated the invention of knife-shaped tools (Ambiru 2010), which could be evidence of foraging groups migrating to Kyushu from the Korean peninsula (C. Bae 2017, in this issue; Matsufuji 1987).

figureFigure 3. Examples of the major stone tools from the early Upper Paleolithic assemblages. 1–6 = trapezoids; 7 = a flake core with small flake scars served for blanks of trapezoids; 8–11 = knife-shaped tools (backed blades); 12–13 = edge-ground axes. Tools 1–3 and 8 are from Jizoden (Kanda 2011). Tools 4–7, 12, and 13 are from Hinatabayashi B (Tani 2000). Tools 9 and 10 are from Happusan (Suto 1999). Tool 11 is from Nawateshita (Yoshikawa 2006).

Chronometric dates, mostly radiocarbon dates based on associated charcoal, demonstrate that the lithic industry characterized by a composite of trapezoids, knife-shaped tools, and edge-ground axes appeared in Japan at 40,000 to 38,000 years ago (Izuho and Kaifu 2015; Tsutsumi 2012; Yamaoka 2010). A substantial number of EUP assemblages (∼500) dated to 38,000 to 30,000 years ago further indicate that modern H. sapiens migrated into the Japanese Archipelago around 40,000 years ago, bringing the new lithic technological complex (Izuho and Kaifu 2015). Culture-chronological division between the Early and Late Paleolithic to characterize lithic industries in East Asia (Gao and Norton 2002; Ikawa-Smith 1978; Seong and Bae 2016) may also be validly applicable to the current Japanese Paleolithic record, although it is necessary to address the question of whether there was Paleolithic human occupation before 40 ka and, if there was, how the earlier Paleolithic record is related to other East Asian Paleolithic records, notably those in China and Korea.

The possibility of an archaeological record before 40 ka was largely dismissed when the Early Paleolithic hoax was exposed in 2000. At that time, it was shown conclusively that the Early and Middle Paleolithic stone tool industries from Miyagi Prefecture were all faked by an amateur archaeologist beginning in the 1980s (Nakazawa 2010; Yamada 2001). Before the fakes were produced, however, the reality of an Early Paleolithic in Japan had been seriously discussed for several sites, such as Sozudai in northern Kyushu and Hoshino in Honshu (Serizawa 1971; Yanagida and Ono 2007). The debate regarding the reality of the Early Paleolithic industry was largely over the issue of whether the fractured flakes were man-made artifacts or not (i.e., geofact). Quaternary geologists suggested that the geological layers of “archaeological artifacts” were derived from alluvial/colluvial sediments that could have created naturally fractured cobbles to make flake-like geofacts (Arai 1971). In contrast, a systematic examination of the angle between the striking platform and the ventral surface of flakes from the pre-40 ka level in the Sozudai site suggested that they were man-made (Bleed 1977), which was largely supported by the proponents of the long-term chronology in the Japanese Paleolithic (Serizawa 1982). Although debate over these sites was shelved while the sensational finds were being “discovered” in Miyagi Prefecture, since the hoax was exposed, many of these sites have subsequently been revisited (e.g., Hagiwara 2006; Ikawa-Smith 2016; Matsufuji 2010; Naruse 2010; Sato 2016; Wada 2016). The candidate assemblages for occupation of the archipelago before the Upper Paleolithic are Kanedori (Tohoku region, NE), Takesa-Nakahara (central Japan, C), Sunabara (southwestern Honshu, SW), and Iriguchi (northern Kyushu, SW; see fig. 2). Multiple criteria have been employed to assign them to before the Upper Paleolithic. First, flaking and retouch technologies have been used. Besides mechanical criteria to distinguish flakes from geofacts (Barnes 1939; Bleed 1977), a peculiar flaking technology called “obtuse angle technology” (Nagai 2011) that is often found in spheroids in the “Lower/Middle Paleolithic” industry in South Korea (Lee 2015) is chosen. The second criterion is whether these stone tools are really different from or similar to the earliest Upper Paleolithic assemblages (i.e., edge-ground axes, knife-shaped tools, and trapezoids) with respect to patterns in tool morphology, reductive technology, and raw material use (e.g., Matsufuji 2010; Suto 2006). For example, Matsufuji (2010:196) suggests that crude and large tools with two small flakes recovered from the Kanedori IV layer are different from the EUP industry, and therefore it is attributed to the “broader East Asian core and flake tool tradition.” The third criterion is the chronological age of the assemblage. Instead of using chronometric dates associated with tool assemblages from before the Upper Paleolithic, Japanese Paleolithic archaeologists have usually employed tephras to develop culture-stratigraphic sequences.

Based on the above multiple criteria, most of the Japanese assemblages from before the Upper Paleolithic cannot support the arrival of humans before 40 ka. However, some recently excavated sites, notably Sunabara, have been investigated through examination of site integrity (Matsufuji and Uemine 2013; Uemine, Matsufuji, and Shibata 2016) and microscopic analysis of fracture mechanics in rhyolite (a coarse-grained material recovered from the site) to identify the man-made nature of lithic artifacts (Uemine 2014). These efforts may eventually stand up to further scientific scrutiny to support an MIS 5e human arrival in the archipelago, as some researchers propose. However, in a case like Sunabara, researchers will be further required to explain how man-made “artifacts,” naturally fractured debris, and naturally transported pebbles were all recovered together in the same alluvial sediments (i.e., layer VIa; Uemine 2014). Only a thorough analysis of the site formation processes may really answer this question.

Among the other artifacts, the lithic assemblages from Kanedori layers IV and III are the most promising lithic artifacts attributable to before the Upper Paleolithic in Japan (Kuroda et al. 2005, 2016; Matsufuji 2010). The lower level of Kanedori layer IVb, where the lower assemblage was recovered, has multiple tephras that were secondarily deposited, suggesting that the age of layer IVb is in the time range of 50,000 to 90,000 years ago (Soda 2005; Yagi 2005). Despite the seeming credibility of stratigraphy, lithic artifacts, and tephra-assisted chronometric dates in the Kanedori assemblages from before the Upper Paleolithic, the number of candidates for Japanese lithic assemblages from before the Upper Paleolithic is still small. Even among the 16 so-called assemblages, there are surface collections (e.g., Kaseizawa) that are undateable (Sato 2016:31, table 1). More detailed evaluation of the characteristics and variability in those assemblages requires further systematic comparison through technological and morphological studies (e.g., Bleed 1977; Nagai 2011). Given the sporadic and sparse occurrence of those candidates for sites from before the Upper Paleolithic, categorizing them into the notion of the “Early Paleolithic” and the extent to which they are comparable to the archaeological record in the East Asian mainland (e.g., Gao 2013; Gao and Norton 2002; Wang 2005) will be an important future research avenue.

Discussion

How much do we know about the Pleistocene human population history in Japan, and how much do we not know? With respect to human migratory routes into the Japanese Archipelago, of the six hypothesized routes of human entry (fig. 1), the routes from the Korean Peninsula and southern China to Kyushu (i.e., routes 1 and 5), a southern part of Paleo-Honshu Island, are the most parsimonious based on the Paleolithic site density and technological and morphological comparisons of formal stone tools between Japan and Korea during the EUP. This route was likely, given that hunter-gatherer population density in the adjacent regions would have been higher than Paleo-Honshu at the time of the earliest population entry. For example, researchers have identified an increasing number of Middle and Late Pleistocene sites in southern China (e.g., Pei et al. 2013; Shen and Keates 2003; Wang 2003, 2005; Wei et al. 2017), suggesting that the Late Pleistocene hunter-gatherer population density in southern China was higher than that in Paleo-Honshu. In contrast with the extensive Paleolithic record in Japan, the Pleistocene human fossil record is primarily concentrated in the Ryukyus. This implies that Upper Paleolithic hunter-gatherers had already migrated into the far southern Japanese islands by seafaring, although the migratory route to get to the Ryukyus is still not clear. It is possible that the initial foragers to arrive in the Ryukyus came from Taiwan in the south (Kaifu et al. 2015) or from southern Kyushu in the north. The latter route was present at least during the Holocene (Obata, Morimoto, and Kakubuchi 2010; Yamazaki 2012).

To further complicate the various migration models, ancient DNA data largely support gene flow from eastern Siberia to Hokkaido, possibly since the LGM. If this were the case, Pleistocene population dynamics were more complex than the admixture model, which assumes population continuity from the Paleolithic to Jomon, followed by the admixture of late Holocene Yayoi peoples, as outlined in the dual-structure model (Hanihara 1991).A more complex picture of Pleistocene hunter-gatherer migrations into the Japanese Archipelago is legitimately implied from the long- and short-term chronologies of the Japanese Paleolithic record. In the framework of the long-term chronology, the question is the extent to which human populations before the Upper Paleolithic (>40 ka) contributed to the establishment of subsequent hunter-gatherer populations since 40 ka. Regardless of the relationship between populations, given the scarce evidence of credible human occupations before 40 ka, which has so far only been provided from a small number of archaeological sites (e.g., lithic industry from the Kanedori before 50 ka), the human population size before 40 ka in Japan was smaller than that of the Upper Paleolithic. In stone tool technology, although it could be an effect of small sample size (n = 40), there seems to have been a change from the unstandardized retouched tools and heavy-duty tools in the industry before 40 ka, as represented by the Kanedori III assemblage, to the formal and standardized stone tool inventory consisting of trapezoids, knife-shaped tools, and edge-ground hand axes in the EUP. This change further suggests that the EUP tool inventory and technology were independently invented among hunter-gatherers before the Upper Paleolithic. In contrast, the currently dominant short-term Paleolithic chronology may pose a different explanation for technological change. In the short-term chronology, the EUP hunter-gatherers were the first population to enter into the Japanese Archipelago. In this context, the EUP tool inventory and blade technology were all brought into Japan, and the subsequent proliferation was the result of relatively rapid population expansion across the archipelago (i.e., demic expansion; Cavalli-Sforza, Menozzi, and Piazza 1993). The Upper Paleolithic demic expansion in Japan syncs well with the modern Homo sapiens single-dispersal model out of Africa and rapid dispersal into South Asia (e.g., Forster and Matsumura 2006; Mellars 2006a, 2006b). However, Upper Paleolithic lithic industries that appear after the end of the EUP (∼30,000 years ago) exhibit extensive regional variation, particularly in the technological, morphological, and stylistic characteristics of the complexes represented by the knife-shaped tools (e.g., Morisaki 2012; Ohyi 1968; Yoshikawa 2010), bifacial points (e.g., Hashizume 2015), and microblade cores (e.g., Sato and Tsutsumi 2007). The observed variation might have been created by a combination of human migrations from the East Asian mainland and endemic technological invention and transformation among Upper Paleolithic hunter-gatherers between the different microregions in the Japanese Archipelago. The interactions and foraging across the boundaries of microregions are often perceived in archaeological patterns, including the long-distance transportation of obsidian (e.g., Tsutsumi 2010) and isolated occurrences of regionally stylistic weapons outside of their core areas, such as the Kou-type knife-shaped tools (e.g., Kato 1975; Morisaki 2012). Moving forward, it will be critical to evaluate the extent to which indigenous hunter-gatherer population density at the microregional scale and the size of populations dispersing from the East Asian mainland covaried and influenced cultural change and variation on the archipelago. Given the complex nature of the Paleolithic archaeological record, human occupation history in Japan is likely compatible with a multiple-dispersal model of H. sapiens (e.g., Bae and Bae 2012; Boivin et al. 2013; Lahr and Foley 1994; Petraglia et al. 2010).What makes the population history in Japan complicated is that the migration from the north via Paleo-SHK was significantly later than for Paleo-Honshu. While a small number of trapezoids that are morphologically comparable to those of the EUP in Paleo-Honshu have been identified in some assemblages in eastern Hokkaido, allowing some archaeologists to place them in late MIS 3 (e.g., Izuho and Takahashi 2005; Oda and Morisaki 2016), archaeological assemblages from the sites having secure associations of chronometric dates and stratigraphy in Hokkaido only appear at the onset of the LGM, 25,000 years ago (Izuho et al. 2012). Flake technology, blade, and microblade technologies were incorporated into the LGM technocomplex in Hokkaido (Izuho et al. 2012; Nakazawa and Izuho 2006, Nakazawa et al. 2005), which later converged into the microblade technocomplexes (Nakazawa and Yamada 2015). This development likely resulted from a combination of independent innovation, cultural transmission, and demic expansion from eastern Siberia and Paleo-Honshu in and after the LGM (e.g., Buvit et al. 2016; Graf 2009; Nakazawa et al. 2005; Nakazawa and Yamada 2015). Why the initial occupation of Paleo-SHK lagged behind that of Paleo-Honshu by some 15,000 years is another area that needs to be further investigated.Although the number of migratory events is difficult to tease out from the current archaeology, human paleontology, and human genetic records, it is likely that it was the result of the admixture of two opposite large migratory events, similar to the Korean Upper Paleolithic (K. Bae 2010; Bae and Bae 2012). This population admixture likely occurred during MIS 2 (30,000 to 11,500 years ago) and involved an influx of hunter-gatherers from the north and south. Evident increases in the number of sites and stone tool technological variability during MIS 2 in both Paleo-Honshu and Paleo-SHK (e.g., Nakazawa and Yamada 2015; Ono et al. 2002; Suto 2006) could be explained by demographic increase and an associated cumulative adaptive culture model (e.g., Henrich 2004; Shennan 2001; Powell, Shennan, and Thomas 2009, 2010).An examination of current evidence in Paleolithic archaeology, human paleontology, and human genetics in Japan necessarily provides a complex picture of Late Pleistocene demographic history. In the vast region of Asia, describing the Pleistocene population history in the Japanese Archipelago will doubtlessly be important in understanding human colonization and evolutionary history. Moreover, the accumulated Paleolithic record in Japan has the potential for improving understanding of the complexity of Pleistocene hunter-gatherer cultural and biological evolution.”

A more recently proven “lost culture”, the arrival of the Okhotsk culture from the north whose Siberian Amur region genetic components are significantly admixed with the Ainu, and whose genetic signals can be seen even in northeast Japan — is still relatively unknown today, despite having had a major influence on introducing barley agriculture, bear ceremonies and whaling culture to the northeast parts of Japan. See Barley dispersal patterns mirror the settlement of a forgotten culture from the north 

Finally, if you take into account all of the above recent literature, the above dispersal scenarios are easier to reconcile with recent admixture models and analyses of genetic affinities between both ancient DNA and MtDNA of modern Asians…

… see the analyses of both McColl et al.’s 2018 Ancient genomics reveals four prehistoric migration waves into Southeast Asia as well as Kanzawa-Kiriyama’s 2019 paper, Late Jomon male and female genome sequences from the Funadomari site in Hokkaido, Japan

Large scale whole genome sequencing 2020 study elucidates the genetic and phenotypic landscapes of mtDNA in the Japanese population

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Fig. 1: High-resolution spectra of mtDNA haplogroups in the Japanese population.

From: Genetic and phenotypic landscape of the mitochondrial genome in the Japanese population  (Open Access pub. under Creative Commons license)

c Stacked bar plots of the frequencies of the macrohaplogroups within the geographical regions in Japan and subpopulations from 1KG. The geographical regions in Japan are defined as Hokkaido, Tohoku, Kanto-Koshinetsu, Chubu-Hokuriku, Kinki, Kyushu, and Okinawa from northeast to southwest areas of Japan, as described elsewhere42.

Below is an excerpt of the open access 2020 paper:

Yamamoto, K., Sakaue, S., Matsuda, K. et al. Genetic and phenotypic landscape of the mitochondrial genome in the Japanese population. Commun Biol 3, 104 (2020). https://doi.org/10.1038/s42003-020-0812-9 PDF

Abtstract

The genetic landscape of mitochondrial DNA (mtDNA) has been elusive. By analyzing mtDNA using the whole genome sequence (WGS) of Japanese individuals (n = 1928), we identified 2023 mtDNA variants and high-resolution haplogroups. Frequency spectra of the haplogroups were population-specific and were heterogeneous among geographic regions within Japan. Application of machine learning methods could finely classify the subjects corresponding to the high-digit mtDNA sub-haplogroups. mtDNA had distinct genetic structures from that of nuclear DNA (nDNA), characterized by no distance-dependent linkage disequilibrium decay, sparse tagging of common variants, and the existence of common haplotypes spanning the entire mtDNA. We did not detect any evidence of mtDNA–nDNA (or mtDNA copy number–nDNA) genotype associations. Together with WGS-based mtDNA variant imputation, we conducted a phenome-wide association study of 147,437 Japanese individuals with 99 clinical phenotypes. We observed pleiotropy of mtDNA genetic risk on the five late-onset human complex traits including creatine kinase (P = 1.7 × 10−12).

… Here we provide a comprehensive analysis that characterizes the genetic and phenotypic landscape of mitochondria in the Japanese population. (i) We constructed a high-resolution mtDNA variant list and haplotype classifications in the Japanese population based on deep WGS of ~2000 individuals. (ii) We conducted unsupervised clustering of the mtDNA variant patterns and assessed their correspondence with defined haplogroups by using a set of machine learning (ML) methods. (iii) We quantitatively assessed positional distributions and LD structure of the mtDNA variants by parallelly comparing with those of nDNA. (iv) We performed genome-wide scans to investigate the mtDNA–nDNA and mtCN–nDNA genotype associations. (v) Finally, we conducted mitochondrial genomewide genotype imputation of genome-wide association study (GWAS) data of more than 140,000 Japanese individuals, by using the population-specific WGS reference data. We then conducted a PheWAS [Phenome-wide association study] of 99 complex human disease and quantitative traits.

RESULTS

High-resolution mtDNA variant and haplogroup lists in the Japanese population

We re-analyzed the previously reported WGS data of the Japanese population (n = 1928)18. We realigned the WGS reads on the human reference genome GRCh37, which includes the revised Cambridge Reference Sequence (rCRS, NC_012920.1) as the human reference mitochondrial genome. The rCRS has been widely used in mitochondrial genome analyses including Japanese individuals. We only used the reads uniquely mapped on the mitochondrial region to avoid the contamination of nuclear copies of the mitochondrial genome (nuMTs)19. In this study, we focused on the analyses of homoplasmy. Then, we identified 2023 variant sites, of which 63 sites were multiallelic (the mean depth = 1488; Supplementary Data 1). Of these, 516 variants (25.5%) were newly identified in our WGS data. Minor allele frequency (MAF) spectra indicated that the majority of the identified variants were rare in Japanese; rare variants (MAF < 0.5%), low-frequency variants (0.5% ≤ MAF < 5%), and common variants (MAF ≥ 5%) accounted for 79.3%, 16.4%, and 4.3%, respectively (Supplementary Fig. 1). We observed clear concordances of the alternative allele frequencies with those in the previously reported two Japanese databases (1507 and 1025 variants with 3.5 KJPN [n = 3552] and Giib-JST mtSNP [n = 672], respectively; Supplementary Fig. 2)7,20,21. As previously reported22,23, mutational spectrum indicated a high transition to transversion (Ti/Tv) ratio of 16.44 (Supplementary Fig. 3).

Next, each individual was classified into the mtDNA haplogroup based on a variant list detected by the WGS using HaploGrep (v2.1.14)24. Haplogroups are classifications of the mtDNA haplotypes defined according to a set of the specific mtDNA variants. As mtDNA is a haploid genome, the detected variants could be directly used for haplogroup classification without phasing. Nomenclature of each haplogroup is hierarchically defined based on the number of the letters (from one to nine), which was divided into sub-haplogroups (e.g, “D4b” as three letters). The number of the haplogroups monotonically increased from the macrohaplogroup (n = 11 at one letter) to the sub-haplogroups with larger number of letters (n = 310 at nine letters; Supplementary Data 2). Increments in the number of the haplogroups became limited from seven to nine letters (Fig. 1a, b).

The frequency distribution of each haplogroup was obtained according to geographical regions in Japan (as defined by BioBank Japan Project: Hokkaido, Tohoku, Kanto-Koshinetsu, Chubu-Hokuriku, Kinki, Kyushu, and Okinawa from northeast to southwest areas of Japan; Supplementary Fig. 4)18 and all the populations from the 1000 Genomes Project Phase 3 (1KG, n = 2504; Fig. 1c)25. In the Japanese population, macrohaplogroups A, B, D, M, and N had more than 1% of frequencies across all the regions. In the regions from Hokkaido to Kyushu, macrohaplogroup D was most prevalent (>28%), followed by M and B. In contrast, the different spectrum was observed in Okinawa (the islandic region at the most southwest area in Japan), where M and B were more prevalent (37.5% and 25%, respectively) than D (18.8%). Furthermore, although the D4a and D4b haplogroups were prevalent from Hokkaido to Kyushu, the D4c haplogroup was prevalent in Okinawa (Supplementary Fig. 5). Although the haplogroup spectra in 1KG East Asians (EAS) were relatively similar to those in Japanese, R was more enriched in 1KG EAS, and D, G, and M were more frequent in Japanese. 1KG populations other than EAS showed distinct haplogroup patterns from the Japanese population. M, A, H, and L were most prevalent in 1KG South Asians, Americans, Europeans, and Africans (AFR), respectively. Especially, 1KG AFR showed the least diversity within macrohaplogroups, of which African-specific macrohaplogroup of L accounted for >90% of frequencies.

Fig. 1: High-resolution spectra of mtDNA haplogroups in the Japanese population

c Stacked bar plots of the frequencies of the macrohaplogroups within the geographical regions in Japan and subpopulations from 1KG. The geographical regions in Japan are defined as Hokkaido, Tohoku, Kanto-Koshinetsu, Chubu-Hokuriku, Kinki, Kyushu, and Okinawa from northeast to southwest areas of Japan, as described elsewhere42.

Fig. 1: High-resolution spectra of mtDNA haplogroups in the Japanese population.

From: Genetic and phenotypic landscape of the mitochondrial genome in the Japanese population

Unsupervised ML approaches deconvoluted mtDNA classification patterns. To evaluate how the defined haplogroups reflect the mtDNA diversity within a population, we conducted unsupervised clustering of the subjects based on the mtDNA variants and evaluated the concordances with haplogroup assignments. We adopted three unsupervised ML classification approaches of phylogenetic approach, principal component analysis (PCA), and uniform manifold approximation and projection for dimensionality reduction (UMAP). We first constructed the phylogenetic tree of the WGS individuals, which was illustrated as an unrooted tree type (Fig. 2a). The tree branch was mainly divided into the major two lineages at the root base, which were known as the “M” and “N” clusters. Each major lineage was further divided into sub-lineages corresponding to the sub-haplogroups. Next, we applied the linear dimensionality reduction technique of PCA and examined up to the 20 PCs. The explained variances were 12.6% for the top 20 PCs. As the two-dimensional plot of the PC1 and PC2 was difficult to fully capture the cluster classifications (Supplementary Fig. 6), we adopted the three-dimensional plot consisting of the top three PCs (Fig.2b). The major M and N clusters were also illustrated as distinct groups in the PCA plot. In contrast to the M cluster, the N cluster was further divided into sub-haplogroups, such as A5a, B4a, B4b, B4c, B5*, F1a, F1b, N9a, and N9b.

Fig. 2 Unsupervised ML-based sample classification of the Japanese mtDNA variant data. All the three unsupervised ML method classifications were performed on the WGS variant data of the Japanese population (n = 1928). a The unrooted phylogenetic tree using maximum-parsimony method. b The 3D plot of the top three components of PCA. c The plot of the two components of UMAP. Each color and marker represents haplogroups. Distinction between the M and N haplogroup clusters is displayed with dotted lines in each panel.

Fig 2

Fig 2 UMAP

UMAP is a recently developed nonlinear dimensionality reduction algorithm, which has a merit in preserving the topology of the local and global structure26. Although UMAP has been previously applied to high-dimensional biological data such as single-cell RNA sequencing27, here we applied it to the genomic data of the mtDNA variants. An application of UMAP could classify the subjects into >20 clusters, which was concordant with the pre-defined sub-haplogroups with three letters (Fig.2c). UMAP could differentiate the sub-haplogroups belonging to the same macrohaplogroup in more detail (e.g., D4a, D4b1, D4b2,D4c, D4e, and D4g belonging to D). On the other hand, we did not find the clear delineation between the M and N clusters, which was observed in the phylogenetic approach and PCA. Several sub-haplogroups with small sample sizes were clustered closely with other macrohaplogroups (e.g., M7c clustered with D macrohaplogroup). These observations could be the potential limitations of UMAP. As each ML classification method hadunique advantages to classify mtDNA variations, we would propose a value of applying multiple ML methods to comprehensively visualize haplogroup diversity within a population.Distinct characteristics of mtDNA variant structure from nDNA. Due to lack of recombination and higher mutation rate, LD structure and tag variant distribution in mtDNA are considered to be distinct from those in nDNA, whereas the details have been unclear especially in non-Europeans. Thus, we conducted a comprehensive assessment of these features in Japanese by using the WGS data. In addition to calculate the haplotype correlations (r2) between common mtDNA variant pairs (MAF≥5%,n=80), we estimated those obtained from the randomly selected autosomal phased haplotypes with adjustment on variant positional differences (±8.3 kbp). Fractions of the variant pairs with high correlations were relatively smaller in mtDNA than in nDNA (2.4% and 20.2% of the variant pairs with r2≥0.9 in mtDNA and nDNA, respectively; Fig.3a).When we checked distance-dependent decay of LD, we observed clear LD decay in the nDNA variant pairs according to physical distances between the variant pairs, which would reflect LD blocks (R=−0.092,P=1.0 × 10−7, as highlighted with red; Fig.3b). However, mtDNA variant showed no distance-dependent LD decay (R=0.022,P=0.21). Although there existed controversial discussions16,28, our results did not support the hypothesis of potential recombination events in mtDNA. Lack of recombination and relatively weak LD in mtDNA should propose that the common mtDNA variants are sparsely tagged bythe surrounding single-nucleotide polymorphisms (SNPs) when compared with nDNA. The numbers of the tag variants per common variant were smaller in mtDNA than in nDNA, even when the variant distances were adjusted (Fig.3c). As many as 13.8% of the common mtDNA variants did not have any tag variants with r2≥0.5, whereas only 5.0% in nDNA. Systematic visualization of pairwise LD patterns revealed that mtDNA variants did not constitute LD blocks among neighboring variants (Fig.3d). On the other hand, we observed multiple common haplotypes spanning the entire mtDNA (n=8), which might have been the consequence of lack of recombination and no distance-dependent LD decay. Interestingly, mtDNA variants without any tag variants were mostly identified in the D-loop region, one of the non-coding but functional regions in mtDNA. By using WGS data, we highlighted the characteristics of the mtDNA variants28, which were (i) no distance-dependent LD decay (i.e., absence of LD blocks) and (ii) sparse tagging of the common variants, in non-European population. In addition, we could newly detect existence of the common haplotypes spanning the entire mtDNA. No evidence of mtDNA–nDNA genotype association. Mitochondrial function is regulated not only by the genes encoded in mtDNA but also by those in nDNA. As these two typesof genes confer synergistic biological functions6, co-evolutionof the genetic variants embedded within them5, namely “mtDNA–nDNA genotype association”, has been suggested17. Thus, we tested the hypothesis that there might exist preferential transmission between nDNA and mtDNA at a genotype level. To explore footprints of the mtDNA–nDNA genotype association,we conducted a genome-wide scan to assess genotype distribution dependence between mtDNA and nDNA variants using the WGS data. We investigated 86 common mtDNA (MAF≥5%) and the genome-wide 7,124,343 nDNA variants (MAF≥1%), but there was no significant mtDNA–nDNA genotype association when multiple comparisons were considered (P=5.0 × 10−8/86=5.8 × 10−10; Fig.4a). Even when we focused on the variants located within ±10 kbp of the previously defined nDNA mito-chondrial genes (n=1105)29, we still did not detect a significant association. In addition, we investigated the associations by using the imputed GWAS data (n=141,552; see details in the next section). We analyzed the association between the 8 mtDNA(MAF≥5%) and the 7,402,102 nDNA variants (Rsq≥0.7 and MAF≥1%), but no significant signals were detected (P=5.0 ×10−8/8=6.3 × 10−9). We neither observed the enrichment of the association signals in the mitochondrial-related gene variants in nDNA (Fig.4b).

 

DISCUSSION

UMAP is a recently developed nonlinear dimensionality reduction algorithm, which has a merit in preserving the topology of the local and global structure26. Although UMAP has been previously applied to high-dimensional biological data such as single-cell RNA sequencing27, here we applied it to the genomic data of the mtDNA variants. An application of UMAP could classify the subjects into >20 clusters, which was concordant with the pre-defined sub-haplogroups with three letters (Fig. 2c). UMAP could differentiate the sub-haplogroups belonging to the same macrohaplogroup in more detail (e.g., D4a, D4b1, D4b2, D4c, D4e, and D4g belonging to D). On the other hand, we did not find the clear delineation between the M and N clusters, which was observed in the phylogenetic approach and PCA. Several sub-haplogroups with small sample sizes were clustered closely with other macrohaplogroups (e.g., M7c clustered with D macrohaplogroup). These observations could be the potential limitations of UMAP. As each ML classification method had unique advantages to classify mtDNA variations, we would propose a value of applying multiple ML methods to comprehensively visualize haplogroup diversity within a population.

Distinct characteristics of mtDNA variant structure from nDNA
Due to lack of recombination and higher mutation rate, LD structure and tag variant distribution in mtDNA are considered to be distinct from those in nDNA, whereas the details have been unclear especially in non-Europeans. Thus, we conducted a comprehensive assessment of these features in Japanese by using the WGS data. In addition to calculate the haplotype correlations (r2) between common mtDNA variant pairs (MAF ≥ 5%, n = 80), we estimated those obtained from the randomly selected autosomal phased haplotypes with adjustment on variant positional differences (±8.3 kbp). Fractions of the variant pairs with high correlations were relatively smaller in mtDNA than in nDNA (2.4% and 20.2% of the variant pairs with r2 ≥ 0.9 in mtDNA and nDNA, respectively; Fig. 3a). …

Our study demonstrated a successful imputation of the mtDNA variants by using the large-scale WGS as a reference panel. …

In conclusion, through the large-scale WGS and PheWAS of mtDNA, our study could comprehensively elucidate the genetic and phenotypic landscapes of mtDNA in the Japanese population.

A treasure trove of 103 ‘lost’ drawings by Hokusai emerges in the Piasa auction house in Paris

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Preparatory drawing (hanshita-e) for an illustrated book, mounted on card. Sea monster assailing a ship. Ink on paper.
© The Trustees of the British Museum

Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0)

103 recently uncovered drawings by the Japanese painter and printmaker Katsushika Hokusai were added to the British Museum’s collection. The works, which resurfaced for the first time since 1948 in Paris last year, were created in 1829 for an unpublished book titled Great Picture Book of Everything(Banbutsu ehon taizen zu 万物絵本大全図 )

The small-scale works were previously owned by Art Nouveau jeweler Henri Vever, and the museum said in a release that they had been held in a private collection in France before their reemergence in 2019 at Piasa auction house in Paris. AFP reports details about the discovery, see excerpts below:

 

British Museum acquires ‘lost’ drawings of Japan’s Hokusai

London (AFP)

The British Museum said Thursday it had acquired 103 “lost” drawings from the 19th century of Japanese artist Hokusai, whose internationally renowned work includes the iconic “The Great Wave”.

The black-and-white drawings were composed in 1829, when Hokusai was 70, as illustrations for an unpublished book. They resurfaced in Paris in 2019 after they were last publicly recorded at an auction in 1948.

Their subjects range from the religious and historical to the mythological, and include previously unknown features of Hokusai’s oeuvre such as a musing on the origins of human culture in ancient China, the London museum said in a statement.

“These works are a major new re-discovery, expanding considerably our knowledge of the artist’s activities at a key period in his life and work,” honorary research fellow Tim Clark said.

“All 103 pieces are treated with the customary fantasy, invention and brush skill found in Hokusai’s late works and it is wonderful that they can finally be enjoyed by the many lovers of his art worldwide,” he said.

The drawings are available to view online and will feature in a future, free exhibition, the museum said.

Read the rest of the article here… and also more at Artnews’

British Museum Acquires 103 Recently Resurfaced, Rarely Seen Hokusai Drawings

A study of maize weevils from ancient Jomon pottery shows they stored acorns and chestnuts long before rice cultivation

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Image of a maize weevil impression from the surface of a pottery fragment.
CREDIT: Prof. Hiroki Obata

Researchers from Kumamoto University have studied nationwide maize weevil impressions from 10,000 year-old pottery,  from the southern Japanese island of Tanegashima,  to pottery fragments from the Sannai-Maruyama site in northern Aomori., to Tatesaki archaeological site in Hokkaido.

The study comparing the differential in the size of  nationwide samples of weevil impressions in the west from the 20 per cent larger ones in the northeast revealed that pots in the west stored acorns while pots in northeastern climes stored chestnuts.

As chestnuts are not native to Hokkaido,  the presence of the maize weevils is evidence that the Jomon people of Tohoku  from south of Hokkkaido, carried supplies including chestnuts infested by weevils, over the Tsugaru Strait by ship.

 

***

 

See excerpts from the study below:

Hiroki Obata et al, Discovery of the Jomon era maize weevils in Hokkaido, Japan and its mean, Journal of Archaeological Science: Reports (2018). DOI: 10.1016/j.jasrep.2018.10.037 Dec 19, 2018

Ancient Japanese pottery includes an estimated 500 maize weevils
by Kumamoto University.

Researchers have discovered an ancient Japanese pottery vessel from the late Jomon period (4500-3300 BP) with an estimated 500 maize weevils incorporated into its design. The vessel was discovered in February 2016 from ruins in Hokkaido, Japan. This extremely rare discovery provides clues on the cultivation and distribution of chestnuts, food in the Jomon era, and the spirituality of ancient Japanese people.

Maize weevils are beetles of the Dryophthorinae subfamily, and are destructive pests of stored rice and grains. By 2003, Jomon-period pottery and pottery fragments containing foreign-body impressions had been collected by various researchers from multiple archeological sites around Japan. Surveys of these impressions exposed hundreds of seed and insect traces on and in the pottery. Over the years, researchers found that maize weevils constituted over 90 percent of all recorded insect impressions.

In 2010, Professor Obata’s research group from Kumamoto University (KU) in Japan found maize weevil impressions in 10,000 year-old pottery that had been recovered from the southern Japanese island of Tanegashima. They showed that maize weevils, which were thought to have come from the Korean Peninsula, had damaged stored food, such as acorns and chestnuts, long before rice cultivation began in the area.

In 2012, the KU research group found maize weevils impressions in pottery fragments from the Sannai-Maruyama site in the northern Japanese prefecture of Aomori. The fact that weevils inhabited an area with a cold winter is an indicator for the distribution food by humans and a warm indoor environment that persisted throughout winter. It is presumed that weevil infestation of stored food was well underway in the Jomon period.

Interestingly, when comparing the body size of 337 maize weevil impressions found nationwide, the team discovered that the body length of maize weevils from eastern Japan was about 20 percent longer than that of western Japan. It is presumed that this body-length discrepancy is due to the different nutritional values between the types of foods they infested—the sweet chestnuts of eastern Japan vs the acorns of western Japan.

Chestnuts are not native to Hokkaido and previous studies surmised that people carried them to the northern Japanese island. The discovery of weevils at the Tatesaki archaeological site in Hokkaido is evidence that the Jomon people of Tohoku (south of Hokkaido) carried supplies, including chestnuts infested by weevils, over the Tsugaru Strait by ship.

See more photos and read more about the research here.

References and further readings:

Hiroki Obata et al, Discovery of the Jomon era maize weevils in Hokkaido, Japan and its mean, Journal of Archaeological Science: Reports (2018). DOI: 10.1016/j.jasrep.2018.10.037 Dec 19, 2018

Ancient Japanese pottery includes an estimated 500 maize weevils -The rare discovery of a vessel with a high number of weevils provides clues to life in ancient Japan (Kumamoto University website) The images above are used in compliance with the image use restrictions from this webpage. See more photos and read more about the research here.


During which season did the Jomon people gather and eat shellfish?

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An exhibit featured this week at Tama Center’s Jomon Village Archaeological Center reveals that the Jomon people collected shellfish mostly during the spring to summer seasons.

Exhibit at Tama Center’s Jomon Village Archaeological Center

After eating the shellfish, the empty shells would be deposited in a dumping site in a location away from the village as can be seen in the left foreground of the replica village below.


Scientists have studied the shells found in the shell mound and determined that they were gathered mostly during the spring to summer period.

Shellfish of Nishigahara She’ll Mound collected by season

How were they able to do this? They were able to do so by studying the growth line of the outer shell layers.

Here is a cross-section of the Nishigahara shellfish mound, which is essentially the remains of a large kitchen dump of the Late to Final Jomon period.

All rights reserved. Text and photos of exhibits by Heritage of Japan.

Maedakochi site: Jomon village life during the Incipient Jomon Period

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Maedakochi site and the recreated landscape of Tama Hills during the Incipient Jomon Period (Photo: Heritage of Japan)

First things first…

Announcement! We are slowly (it will be a laborious work in progress) moving to a new website (still with WordPress), uploading a few pages at a time. If you like the look of the new website, or have other feedback, please leave a comment over there.  Meanwhile, we continue to blog and keep all our previous content here. 

I have begun a series on the Jomon Village Historic Garden of the Tama Hills (which I have visited probably over 10 times!). Archaeologists have excavated 770 out of the 964 sites from the Tama Hills (aka the Tama New Town to locals). A “Jomon village” based on settlement traces was recreated, with archaeological exhibits from 40 years of diggings on display at the adjacent Exhibition Hall run by the Tokyo Metropolitan Archaeological Center.

Archaeological sites of the Tama Hills which straddles the cities of Tama Machida, Inagi and Hachioji today. The red star is the location of the Jomon Village / Tokyo Metropolitan Archaeological Center’s Exhibition hall

People began living in this area around 32,000 years ago, towards the end of the Ice Age when the climate was colder and more like Hokkaido’s, and when people hunted game such as Giant Elk and Naumann elephants, salmon from the rivers and large animals that lived in the coniferous forests. (They left behind stone tools and burnt stones from their fires, but I will post in another episode about the Upper Palaeolithic traces and remains, because the focus of this post is the Jomon people from the Maedakochi site).

The Maedakochi site (see photo at the top of the page) is currently the central exhibit being featured at the Exhibition Hall of the Jomon Village.

The discovery of the Maedakochi site is an important one as it is one of the earliest sites of the Jomon people belonging to the Incipient Jomon Period radiocarbon dated to 15,500 years ago. Being at the transition phase between the Upper Paleolithic and the Neolithic Jomon Period, the excavation of the site reveals important clues to neolithic lifestyles, such as why early pottery was invented and what the ceramic pots contained and were used for. Numerous salmon teeth and bones were found at the site too. Much recent literature has emerged based on analysis of the archaeological evidence from this site, indicating that the Jomon neolithic package likely was part of an entire Aquatic Neolithic complex extending to Sakhalin Island and to the Russian Far East Amur region. To know more, read the rest at The earliest Jomon developed and used pottery for fishing salmon and molluscs 15,500 years ago

Migrants arrive to form the Okhotsk culture in northern Hokkaido, bringing a type of barley different from that of the Satsumon people

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Here, I’d like to highlight data from two papers about the prehistoric dispersal routes of barley from Eurasia into Hokkaido.  From the first, Lister 2018, I draw our attention to the proposed routes by which mainly six distinctly different types (and distributions) of domesticated barley arrived from the west to Eastern Eurasia into the terminal end in Japan.  Note in particular the two different routes of barley into Japan:  The northern route into Hokkaido via Sakhalin Island vs. the route from the Chinese coast to southwestern Japan in Lister’s map below.


Proposed routes of spread of six types of domesticated barley (vulgare) genepools  Source: Lister, 2018

The second paper by Leipe et al, see the excerpted analysis that follows below considers new evidence about the little known Okhotsk culture, who were migrants who came from the north through Sakhalin Island (likely from the Lower Amur/Russian Far East region) to Hokkaido, between 600~1000 AD. While the Okhotsk people were known to have intensively exploited marine resources, this study considers new evidence that they supplemented their marine diet with a far wider range of foods than marine resources including terrestrial mammals such as deer, fox, rabbit, and marten, domesticated pigs and dogs as well as edible wild plants including Aralia (spikenard), Polygonum(knotweeds), Actinidia (Chinese gooseberry), Vitis(grapevines), Sambucus (elderberry), Empetrum nigrum(crowberry), Rubus sp. (blackberry), Phellodendron amurense (Amur corktree), and Juglans (walnut). In addition to broomcorn millet, foxtail millet, the paper considers whether the evidence of the new type of barley grains that they brought with them could have been cultivated for agriculture, for ritual use, or on a lighter scale.

This paper is significant as it extends our knowledge beyond what is known about the intensive marine resources subsistence strategy of the Okhotsk culture, based on its study of botanical samples from the Hamanaka site on Rebun Island. In particular, researchers observed that there are two distinctive phenotypes of barley grown in Hokkaido and that clear boundaries are observed of cultivation between the Satsumon hulled barley in the south and the naked barley grown by the Okhotsk culture. Consequently, a picture emerges on the different origins and interactions of the Okhotsk vs Satsumon cultures: the cultivation by the Okhotsk culture of naked barley over 500 years, and the arrival and expansion of the Satsumon with hulled barley towards the 10th ~11th centuries, reflect the pattern of interactions between the two cultures, and as well as evidence of the timing of the arrival and expansion of the Yayoi people into Hokkaido (click to see Leipe et al.’s Fig 1 map).

The 2017 paper by Leipe et al., Barley (Hordeum vulgare) in the Okhotsk culture (5th–10th century AD) of northern Japan and the role of cultivated plants in hunter–gatherer economies

This paper discusses archaeobotanical remains of  naked barley recovered from the Okhotsk cultural layers of the Hamanaka 2 archaeological site on Rebun Island, northern Japan. Calibrated ages (68% confidence interval) of the directly dated barley remains suggest that the crop was used at the site ca. 440–890 cal yr AD. Together with the finds from the Oumu site (north-eastern Hokkaido Island), the recovered seed assemblage marks the oldest well-documented evidence for the use of barley in the Hokkaido Region. The archaeo-botanical data together with the results of a detailed pollen analysis of contemporaneous sediment layers from the bottom of nearby Lake Kushu point to low-level food production, including cultivation of barley and possible management of wild plants that complemented a wide range of foods derived from hunting, fishing, and gathering. This qualifies the people of the Okhotsk culture as one element of the long-term and spatially broader Holocene hunter–gatherer cultural complex (including also Jomon, Epi-Jomon, Satsumon, and Ainu cultures) of the Japanese archipelago, which may be placed somewhere between the traditionally accepted boundaries between foraging and agriculture. To our knowledge, the archaeobotanical assemblages from the Hokkaido Okhotsk culture sites highlight the north-eastern limit of prehistoric barley dispersal.
Seed morphological characteristics identify two different barley phenotypes in the Hokkaido Region. One compact type (naked barley) associated with the Okhotsk culture and a less compact type (hulled barley) associated with Early–Middle Satsumon culture sites. This supports earlier suggestions that the “Satsumon type” barley was likely propagated by the expansion of the Yayoi culture via south-western Japan, while the “Okhotsk type” spread from the continental Russian Far East region, across the Sea of Japan. After the two phenotypes were independently introduced to Hokkaido, the boundary between both barley domains possibly existed ca. 600–1000 cal yr AD across the island region. Despite a large body of studies and numerous theoretical and conceptual debates, the question of how to differentiate between hunter–gatherer and farming economies persists reflecting the wide range of dynamic subsistence strategies used by humans through the Holocene. Our current study contributes to the ongoing discussion of this important issue...
People in northern Japan, similar to those in Greenland, Arctic regions of Asia, and the American West Coast, remained “complex” hunter–fisher–gatherer well into the historic period [19]. Local Jomon populations of Hokkaido continued a foraging lifestyle [20] until the middle of the 1st millennium AD when they were replaced[?] by Okhotsk cultural communities in the north and by Satsumon cultural communities in the central and the southern parts of the island [21]. Both of the latter cultures are commonly identified as hunter–fisher–gatherers [19]; however, their archaeological remains show evidence for the use of metals and the cultivation of crops [8]. The extent of their productive economy has not been fully studied. To date, the only clear evidence for the cultivation of barely and other crops by Satsumon people in the late 1st millennium AD comes from a single excavation site, which is located in the municipality of Sapporo [8]. There are more data showing that their contemporary neighbours to the north, the Okhotsk culture in Hokkaido, were cultivating both broomcorn (Panicum miliaceum) and foxtail (Setaria italica) millet and barley [8]. However, there have only been a few archaeobotanical studies on Okhotsk sites in Hokkaido and they are entirely published in local Japanese periodicals (e.g. [22, 23]), which are often not available to the international scientific community….
The Okhotsk Culture
The people of the Okhotsk archaeological culture are regarded as a hunter–gatherer society with an economy that strongly relied on marine resources. They occupied a widespread maritime environment, mainly along the southern and eastern littoral margins of the Sea of Okhotsk including northern and north-eastern Hokkaido (see Fig 1C for archaeological site distribution), Sakhalin Island, and the Kurils (Fig 1A). In the Hokkaido Region, the peak of the Okhotsk cultural occupation dates from the 6th to the 8th century AD (see [26] and references therein). Based on pottery style, the “Okhotsk cultural sequence” in northern Hokkaido is divided into three chronological stages comprising the (1) Susuya culture (2nd–5th century AD), which is often referred to as incipient or Proto-Okhotsk, (2) the Towada, Kokumon, Chinsenmon, Haritsukemon, and Somenmon cultures (6th–8th century AD) regarded as the main stages, and (3) the Motochi culture (9th–10th century AD) as the final stage [27]. While Okhotsk cultural traits persisted through the Tobinitai period in eastern Hokkaido until the 12th century AD, replacement or assimilation of the Okhotsk culture in northern Hokkaido by Satsumon/Proto-Ainu populations originating from the central and southern areas of Hokkaido was completed by the end of the 10th century AD …

Archaeologists believe that the Okhotsk culture people migrated to Hokkaido from the north (i.e. Sakhalin Island), first occupying Rebun and Rishiri islands as well as the northern tip of Hokkaido and subsequently dispersing eastwards along the coast of the Sea of Okhotsk [28]. Results of archaeological and genetic studies suggest that the Okhotsk population probably originated from the lower Amur River basin (e.g. [2931]). The population spread onto the islands bordering the Sea of Okhotsk, which is believed to have been due to socio-political conflict [31]. There is also evidence for the onset of cooler climatic conditions in the lower Amur River basin around the end of the 1st millennium BC [32, 33]. These climate changes may have played a role in the southward spread (ca. 500 AD) of these people to Hokkaido [34] and their later absorption/replacement (by ca. 1000–1200 cal yr AD; [26, 27]).

A defining trait of the Okhotsk culture is its subsistence strategy, traditionally thought to be a specialised system of marine resource extraction [26, 35]. This is reflected by the geographic distribution of sites along coastal regions (Fig 1C) and confirmed by archaeological studies of faunal remains and tool assemblages, which indicate intensive marine hunting, fishing, and gathering activities (e.g. [31, 36, 37]). Nitrogen stable isotope studies on human remains also point to a diet with high proportions of protein derived from marine organisms (e.g. [35, 38, 39]). Analysis of human bone collagen revealed a relative contribution of marine protein in the range of 60–94% for individuals from Rebun Island and 80–90% for individuals from eastern Hokkaido [39]. However, there is enough evidence to suggest that the diet of the Okhotsk people may have been much more diverse than the isotopic data imply. People likely supplemented the maritime resources with terrestrial mammals such as deer, fox, rabbit, and marten [37]. Cut marks on bones from domesticated dogs suggest that they were also part of the diet [36], and remains of domestic pigs are limited to northern Hokkaido [26]. In addition, there is evidence for the use of edible wild plants including Aralia (spikenard), Polygonum(knotweeds), Actinidia (Chinese gooseberry), Vitis(grapevines), Sambucus (elderberry), Empetrum nigrum(crowberry), Rubus sp. (blackberry), Phellodendron amurense (Amur corktree), and Juglans (walnut). Furthermore, as already noted, broomcorn millet, foxtail millet, and barley grains have been recovered from sites in this cultural horizon (see [26] and references therein). Admittedly, we know very little about the role of any of these plants in the economy, or whether the crops had a dietary or ritual function. ….

Within the RFE, 15 sites are situated in southern Primor’e (Fig 1B, no. 2–16) and one in the western Amur River valley (Fig 1A, no. 1). The Primor’e sites are associated with early Iron Age Yankovskaia (ca. 850–350 cal yr BC) and Krounovskaia (ca. 500 cal yr BC–200/300 cal yr AD) cultures, the Iron Age Ol’ginskaia culture (ca. 300 cal yr BC–300/400 cal yr AD), the early medieval Mohe culture (ca. 5th–11th century AD), the Bohai State (698–926 cal yr AD), the period following the defeat of the Bohai State (10th century AD), and the Eastern Xia State (1215–1233 AD). The barley finds from the western Amur River valley site represent Troitskii variant of the Mohe culture (end of 8th–9th century AD). From these sites there are in total 25 barley records of which 17 contain measurement data for individual grains. The number of grains per record ranges from 1 to 40. … A total of eleven samples of archaeobotanical barley from south of the Okhotsk culture domain were also considered in this study. They originated in the Hokkaido Region from Early to Middle Satsumon culture (Fig 1C, no. 17–24) and northern Tohoku Heian period (Fig 1C, no. 25–27) sites and date to 8–10th and 9–11th centuries AD, respectively. Scholars have suggested that during the Okhotsk Tobinitai stage (11th–12th century AD; [26]), which emerged in the north-eastern part of Hokkaido, there were enhanced interactions with Late (ca. 1000–1200 cal yr AD) Satsumon groups [54]. Therefore, the review of Satsumon barley is limited to the early and middle stages. The information available for the Early and Middle Satsumon sites (n = 8) is restricted to mean values for L/W ratios, which vary between 1.65 and 2.58 with an average of 2.16. Absolute numbers of measured seeds are not provided in the publications, except for one site (Fig 1C, no. 24). Information from northern Tohoku is based on measurements of three assemblages containing 2, 7, and 50 barley specimens. Their length to width ratios equate 2.3, and their median is 2.29.

For Early and Middle Satsumon barley assemblages information on grain shape (i.e. L/W ratio) is only available in the form of arithmetic means, a statistic which is sensitive to outliers. However, these values are, with one exception, all within the inner whiskers of box-plots of the L/W values for barley from northern Tohoku sites (Fig 4), thus regarded as representing comparable barley varieties. One notable exception is the relatively low mean value of 1.65. Given its small population of five grains and strong offset to the other means, this sample may be regarded as an outlier. The results delineate that the charred barley seeds recovered from Okhotsk culture sites are the most compact. Although still plumper, they are more comparable in shape to grains from the RFE than to those found in Satsumon sites on Hokkaido and Heian period sites in northern Tohoku. While the barley from south of the Okhotsk domain appears to be longer and narrower, the Okhotsk culture and RFE varieties are more compact. The differences in L/W ratio indicate that the long grains comprise hulled and the compact ones naked barley. The ‘nud‘ allele for naked barley is monophyletic [55], thus genetically distinct from hulled barley. This implies a sharp difference between the barley that was grown in these different regions.

Discussion

There is evidence that human migrations from the north have played an important role in the prehistory of Hokkaido and other parts of the Japanese archipelago. This includes the intrusion of Siberian Palaeolithic hunter–gatherer groups around the Late Glacial Maximum (ca. 20,000 cal yr BP; [60]) and immigration ca. 15,000 cal yr BP, with the latter introducing microblade technologies on Hokkaido and Honshu [61, 62]. While they have not been taken into account for a while (e.g. [63]), recent anthropological studies (e.g. [64]) stress the role of migration from northern regions via Hokkaido also in view of the origins of the Neolithic Jomon culture. The most recent southward movement of prehistoric populations into the northern and north-eastern coastal areas of Hokkaido was that of the Okhotsk culture around the middle of the 1st millennium AD [26]. Though, the Okhotsk groups inhabited a large area along the southern and eastern margins of the Sea of Okhotsk, most of our current knowledge has been derived from archaeological materials recovered in the Hokkaido Region. The archaeobotanical record from the Hamanaka 2 site presented in this study allows for greater insight into the use of plants by the Okhotsk people on Rebun Island (Fig 1D). Calibrated ages (95% confidence interval) of directly dated barley remains from five archaeological layers (IIIa–e) suggest that the crop was used at the site between 430–960 cal yr AD (Table 2) or at a 68% confidence interval between ca. 440 and 890 cal yr AD. This time period roughly corresponds to the late Susuya through mid-Motochi stages spanning between the 5th and 10th century AD [27], thus covering the Okhotsk culture settlement phase in northern Hokkaido as indicated by previous archaeological studies. Given the age of the oldest barley seed F2014-037-003 (440–600 cal yr AD, 68% confidence interval; Table 2), the Hamanaka 2 layer IIIe, together with the single dated grain (428–573 cal yr AD, 95% confidence interval; [65]; S3 Table) from the Oumu site (no. 29 in Fig 1C) represents, the earliest well-documented record of domesticated barley in the Hokkaido Region. The only carbonised barley grain recovered in Hokkaido was collected from the Epi-Jomon level of the K135–4 Chome site within the city of Sapporo [66]. This single barley seed has not been directly dated and its proposed age of ca. 200–400 cal yr AD should be viewed with caution.

The presence of barely at the Hamanaka 2 site appears contemporaneous with a phase of enhanced human-induced vegetation disturbance on Rebun Island as indicated by the pollen record from Lake Kushu (Fig 5). During this time (with a maximum ca. 550–800 cal yr AD), the pollen record shows a decrease in the abundance of arboreal pollen, suggesting deforestation and greater openness of the landscape compared to the preceding and subsequent periods, which are more or less coeval with the Epi-Jomon and Proto-Okhotsk (Susuya) cultures (ca. 100 cal yr BC–500 cal yr AD) and Proto-Ainu culture (ca. 950–1600 cal yr AD), respectively ([26, 27]; Fig 5). The results of the local vegetation reconstruction clearly indicate enhanced human activities on Rebun Island during the main phase of the Okhotsk presence there. On the other hand, reduced impact is evidenced during the Epi-Jomon phase and the time of cultural shifts towards the Classic Ainu period, which may be explained by reduced population size and/or a different pattern of resource exploitation. Regarding the Epi-Jomon, this would conform to identified traits like short-term occupations, high mobility, and low complexity [66, 67]. Ohyi [28] suggests that by the time of the disappearance of the Okhotsk culture at the end of the Motochi stage, the Satsumon people spread into northern Hokkaido, including Rebun Island and neighbouring Rishiri Island. It appears, at least for Rebun Island, that these Satsumon groups weakly impacted the island’s vegetation, which was leading to the recovery of the local fir forests. … Rebun Island is well-known for its Okhotsk culture sites, representative for the northern Hokkaido domain. Here, the presence of the Okhotsk groups continued into the Motochi stage (9th–10th century AD) at a time when the Okhotsk sites in northern Hokkaido became abandoned [28]. However, on Rebun Island the presently discussed Hamanaka 2 assemblage contains the only barley thus far recovered (T. Amano, personal communication). This might be due to a lack of systematic sampling and water flotation at other sites on the island. We identified the recorded barley as naked barley, which is far more commonly found than hulled barley in East Asia [68]. Barley was consumed at the site over a ca. 500-year period throughout the main stages of the Okhotsk culture (Fig 5). Barley was significant and had a long-term role in diet during the peak of the Okhotsk culture in the region. The use of barley is also evident at other sites in north-eastern Hokkaido (Fig 1C), being assigned to the late phase (8th–9th century AD) of the Okhotsk culture [69]. In addition, remains of foxtail and broomcorn millet are reported from several excavations [69]. Japanese palaeobotanists have argued that these crops were used for ritual purposes (e.g. [22]); however, this is hard to defend seeing that they appear in so many domestic contexts across such a large time period. The grains likely supplemented a mixed economic system that relied heavily on wild coastal resources. Although, an alternative hypothesis is that these crops were used to produce alcohol…

[A diverse subsistence strategy]

One reason why some scholars have been hesitant to accept that barley and other cereals were dietary supplements may be that the Okhotsk are generally regarded as a specialised hunter–gatherer culture with a subsistence strongly focusing on maritime food resources. This traditional view of a coastal foraging society has been bolstered by recent human bone isotope studies (e.g. [35, 38, 39]), which revealed a high proportion of absorbed protein derived from marine resources of up to 94% and 90% in northern and eastern Hokkaido, respectively [39]. However, our findings together with results of previous studies illustrate that the Okhotsk relied on a wide range of natural and domesticated foods. Besides the suggested strong focus on marine collecting, fishing, and mammal hunting, the Okhotsk people appear to have employed a broad spectrum of wild terrestrial plant fruits and root tubers (this study, Table 3; [26] and references therein), hunted a variety of terrestrial mammals [37], and also maintained domesticated dogs [36] and pigs [26] as part of their food economy. Indications for plant maintenance also comes from the increase in Lysichiton type pollen in the Lake Kushu pollen record (Fig 5) and carbonised Cyperus sp. root tubers in the Hamanaka 2 flotation samples (Table 3). Both taxa represent plants growing in swampy environments around Lake Kushu, which include edible parts and provide nutritious food. It is conceivable that the local Okhotsk people exploited these plants and even maintained their growth and productivity using suitable tools for tilling as found in contemporaneous cultural strata on Rebun Island [71].

Given the combination of foraging, animal husbandry and the use of barley and other cereals over a wide spatio-temporal array, crop cultivation as a supplementary portion of Okhotsk subsistence seems more plausible. This case study further augments existing examples of (“complex”) hunter–gatherers, occupying the “middle ground” which separates hunting–fishing–foraging societies exclusively depending on wild food resources and agriculturalists with a major focus on managing and producing domesticated plants and animals (e.g. [72, 73]). It has been noted that this middle ground territory is highly complex. As Smith [9] puts it, “this territory between hunting–gathering and agriculture is turning out to be surprisingly large and quite diverse; it has also proven to be quite difficult to consistently describe in even the simplest conceptual or developmental terms”. Smith [74] built his concept of ‘low-level food production’ on earlier observations by Braidwood and Howe [75] as well as Flannery [76], all of whom use the term “incipient cultivation” to describe intermediary strategies between foraging and farming. Many other ethnographers and archaeologists have subsequently noted that there is a wide range of diversity in human economic systems; notably, Boserup [77] points out that the range of land-use strategies reflect an equally broad range of human adaptive economies. There have been different approaches to define the middle ground landscape. Following the conceptual framework of Smith [9], who identified low-level food production (<50% annual caloric budget from domesticates) relating to tended wild plants and/or cultivated/managed plants. The Okhotsk culture may be confidently placed somewhere between the traditionally accepted boundaries between foraging and agriculture. The same view is taken by Crawford [8] who explicitly assumes that the Okhotsk people themselves cultivated barley and millet. Given the evidence for dog and pig husbandry and the cultivation or exchange of barley, the Okhotsk epitomises the complexity and diversity of the middle ground economy. …

… the adoption of domesticates by the Okhotsk people, which occurred in relatively recent prehistory, adds particular value as it bridges the gap between the two foci. Another specific feature of the Okhotsk subsistence strategy is the process of adopting already fully domesticated plants that were, by this time, widely used as staple crops in agrarian societies across Eurasia. In the Okhotsk culture, however, the incorporation of barley and millets does not appear to have had significant socio-economic effects. While the subsistence economy continued to be based on foraging, the society remained egalitarian and “group-oriented” [26]. Similar observations are reported from other regions like Japan as well as Island Southeast Asia and Melanesia. The spread of domesticated rice through the latter two regions appears to have started at the beginning of the 1st millennium BC [81]. However, in most regions, rice remained a minor supplementary crop in subsistence systems mainly based on vegeculture (e.g. taro, banana, and sago production) and foraging [82] until the middle of the 1st millennium AD [83]. In Japan, the oldest botanical remains of domesticated broomcorn and foxtail millet, barley, and rice date between the Middle and Late Jomon periods [17], thus may also indicate an early (pre-Yayoi) introduction of domesticated cereal crops from outside the archipelago as minor subsistence supplements not signifying a fundamental change in dietary pattern.

…the assemblages Okhotsk and Satsumon sites in Hokkaido represent the north-eastern edge of prehistoric barley dispersal across Asia. The upper end (600 cal yr AD) of the calibrated age range (68% confidence interval) of the oldest barley seed contained in the dated Hamanaka 2 sample set is coeval with the onset of the Satsumon culture (beginning 7th century AD), which is believed to have arisen from the Tohoku Region (Fig 1A) Yayoi culture populations driven to Hokkaido by expansion of the first Japanese state [8]. Therefore, the most straightforward inference would be that the barley used by the Okhotsk was derived from Satsumon groups spreading into the central and south-western part of Hokkaido. In fact, previous palaeobotanical work points to a different origin that is further emphasised by the Hamanaka 2 barley seed inventory. In previous studies, Japanese scholars claimed to have identified a short and a long barley type at Okhotsk and Satsumon culture sites in the Hokkaido Region, which they assigned to the crop’s naked and hulled form, respectively (see [69] and references therein). Based on this differentiation and seed morphology, Yamada and colleagues (e.g. [22, 23, 69, 90]) have hypothesised that Okhotsk barley originated from neighbouring regions on the Asian mainland. They found that the highly compact (naked barley) specimens extracted from four Okhotsk culture sites (no. 17–20 in Fig 1C, S3 Table) are distinct from the slimmer (hulled) barley (dated to 8–10th century AD) used by Early and Middle Satsumon groups, but similar to grains identified as naked barley found in the early Iron Age to medieval (ca. mid-1st millennium BC–early 13th century AD) sites in southern Primor’e (RFE). Their morphological comparison of barley grains is based on a length/width ratio dataset. Here, we review their approach by supplementing the available datasets, with eleven (partly unpublished) assemblages from the RFE, three records from northern Tohoku, and the measurements of the Hamanaka 2 site barley (S3 Table). Although naked barley is in general more compact than its hulled counterpart, quantitative morphological comparison allows for objective qualification of the recorded archaeological barley and may provide further confirmation for the proposed Okhotsk barley origin. The Okhotsk barley, which appears to be the most compact of all gathered records (Fig 4, S3 Table), in terms of shape is more similar to its counterparts found in both earlier and later (ca. 850 cal yr BC–1000 cal yr AD) sites located along coastal regions across the Sea of Japan (i.e. southern Primor’e, Fig 1C) and in the Amur River basin (Fig 1A), as opposed to grains recovered from contemporary Satsumon culture sites situated in central and southern Hokkaido. Both the Satsumon and Heian grains, which are morphologically similar to each other, appear generally longer and narrower than those used by the Okhotsk culture. This corroborates Crawford’s hypothesis that the Satsumon culture emerged from the Tohoku Yayoi culture [8], which, when being forced to migrate to Hokkaido, brought their barley with them. Alternatively, this similarity may at least suggest cultural interactions between Satsumon populations and communities in Tohoku. The results also corroborate the hypothesis that the Satsumon type barley represents hulled barely and that the naked Okhotsk barley originated in the continental RFE Region. The minor discrepancies in L/W ratios between the naked barley from Okhotsk sites and sites in the RFE (Fig 4) may reflect morphological differences commonly existing among landrace varieties of crops [91] or may be the result of different environmental conditions or irrigation [92]. This means that the barley used by the Okhotsk culture was either derived by exchange with continental populations (e.g. Mohe culture, Bohai State) from across the Sea of Japan or was brought along and cultivated by the Okhotsk culture from their region of origin (i.e. the lower Amur River basin). Unfortunately, no palaeobotanical studies have been conducted in the lower Amur River basin or on Sakhalin Island, which would have allowed us to trace back potential pathways of a southward barley introduction to Hokkaido. Further evidence for the existence of two barley phenotypes in Hokkaido comes from sites, which post-date the main phase of Okhotsk culture. Both types—the compact (naked) barley found at Okhotsk culture sites and the slim (hulled) barley found at Early to Middle Satsumon sites—are represented in archaeobotanical records from this time indicating that Okhotsk type naked barley cultivation/use continued during times of acculturation (i.e. Tobinitai culture) and the Late Satsumon stage [23]. In sum, these findings suggest that naked and hulled barley spread eastward through Asia and were introduced into the Japanese archipelago via different routes. While the area where hulled barley is recovered parallels the distribution of the Yayoi culture (south-western and central Japan) and the Satsumon culture (south-western and central Hokkaido), naked barley possibly propagated from Primor’e and adjacent regions during the Okhotsk culture spread, into Sakhalin Island and northern/north-eastern Hokkaido (Fig 1A). After the two barley phenotypes were independently introduced to Hokkaido, the boundary between both barley domains (Fig 1A) possibly existed for about 400 years across the island region until the beginning of the assimilation/replacement of Okhotsk populations by the Satsumon culture groups (ca. 1000 cal yr AD; [26, 27]).

Further evidence for a close relationship between the Okhotsk culture and Iron Age/medieval populations from the East Asian continent comes from analogies in subsistence economy as described by Sergusheva and Vostretsov [15]. Like the Okhotsk, the Yankovskaia culture (ca. 850–350 cal yr BC; [15]), which inhabited the coastal regions of today’s northern Korea and southern Primor’e, based its subsistence mainly on a wide range of marine resources. Remains of millets (broomcorn and foxtail) and naked barley found at the majority of sites suggest that these domesticates also played a role in diet there. Primor’e Region, in particular, saw a subsequent two-step advance in agricultural practices by the introduction of additional crops including wheat (Triticum aestivum/compactum), hemp (Cannabis sativa), and legumes during the Krounovskaia/Tuanjie culture (ca. 500 BC–200/300 cal yr AD; [15]) and cultigens including hulled barley (Hordeum vulgare), soy beans (Glycine max), and buckwheat (Fagopyrum esculentum) along with the establishment of early states (e.g. Bohai State) after the middle of the 8th century AD. Despite this progress in agriculture, hunting, fishing, and gathering has continuously been an additional part of the diet. In addition, pig and dog breeding is evidenced during the Yankovskaia culture and the Krounovskaia/Tuanjie culture [71, 93, 94]. These subsistence traits place the above mentioned cultures of coastal RFE and northern Korea also in Smith’s middle ground [9], thus somewhere between hunting–fishing–foraging and agriculture. Systematic water flotation has been practiced for over 30 years in this region, providing archaeobotanical remains from numerous Neolithic–Middle Age sites across Primor’e. These data show that during the Yankovskaia culture, millet and barley cultivation was not the main part of the food economy, and was probably not practiced at every archaeological site [15]. Although crop cultivation seems to have been intensified by the Krounovskaia/Tuanjie culture, there is evidence that these groups, likely due to climatic cooling at the end of the 3rd century AD, partly gave up agricultural practices and re-intensified the exploitation of wild resources [95]. This, on the one hand, emphasises that the transformation towards agriculture is not necessarily a unidirectional progression, as it was once regarded, but is a reversible process. On the other hand, it suggests that crops probably had a long-term utility as complementary foods. This might apply to Okhotsk groups, which retained their once-adopted (low-level) agricultural food production as they migrated and adapted to the maritime landscapes of the Sea of Okhotsk.

Conclusions

The archaeobotanical assemblage from Okhotsk cultural layers at the Hamanaka 2 site (northern Rebun Island, Japan) contained charred grains of compact naked barley. Direct radiocarbon dating indicates long-term use of barley at the site over a period of about 500 years. Together with the finds from the Oumu site, the data that we present marks the oldest well-documented evidence for the use of barley in the Hokkaido Region. Due to the broad error ranges of the calibrated radiocarbon dates of the oldest seed remains (428/440–573/600 cal yr AD, 68% confidence interval), more precise ages cannot be defined at this time. However, it is conceivable that the people of the Okhotsk culture were using this crop since they first arrived in the Hokkaido Region (ca. 500 cal yr AD). Accordingly, barley introduction by the Okhotsk culture would pre-date its adoption or introduction by Satsumon populations by at least a century, which may speak against the hypothesis that barley was introduced to northern Hokkaido by the more agrarian south.

The macrobotanical remains of barley are not enough evidence to argue for cultivation at the site, as opposed to the importing of grains from elsewhere. However, axe- and hoe-shaped bone tools found at nearby sites were likely farming implements and do support the possibility of local cultivation. In addition to low-level cultivation, the archaeobotanical data also suggests that wild plant management was conducted by the people of Rebun Island. The pollen record from Lake Kushu indicates significant local vegetation disturbance (i.e. deforestation) concurrent with the barley record at nearby Hamanaka 2. The reconstructed landscape patchiness may point to land clearance for small-scale crop cultivation. In view of the major role of marine food resources indicated by bone isotope studies, it seems likely that cultivated crops were used as supplementary food or for brewing beer. While future studies will clarify the role of barley in the economy in this region, it does seem clear that there was some kind of low-level crop production, which was complemented by wild foods. …

So far, the archaeobotanical assemblages from the Hokkaido Okhotsk culture sites highlight the north-eastern limit of prehistoric barley dispersal. Seed morphological characteristics identified two different barley phenotypes, which were likely independently introduced to the Hokkaido Region. One highly compact type (naked barley) associated with the Okhotsk culture and a less compact type (likely hulled barley) that is evident in Early–Middle Satsumon culture sites. The much more comprehensive dataset presented in this paper supports earlier suggestions that the “Satsumon type” barley was likely propagated by the expansion of the Yayoi culture from south-western Japan towards north-eastern Japan, while the “Okhotsk type” spread from the continental RFE Region, across the Sea of Japan. Although Okhotsk populations may have obtained barley by exchange, there is growing data that suggest that they cultivated naked barley locally, which they introduced directly from their region of origin (i.e. the lower Amur River basin) via Sakhalin. To further verify this hypothesis, additional palaeobotanical studies on materials from archaeological sites in these areas are essential. Nevertheless, based on existing palaeobotanical evidence, we conclude that the Okhotsk culture represents one element of the long-term and spatially broader Holocene hunter–gatherer cultural complex (including also Jomon, Epi-Jomon, Satsumon, and Ainu cultures) of the Japanese archipelago, which may be placed into Smith’s [9] middle ground subsistence strategy. This middle ground domain may chronologically include the groups dating to the Neolithic–Iron Age interval (ca. 3300 cal yr BC–middle 1st millennium AD) and such cultures as Zaisanovskaia, Yankovskaia, and Krounovskaia of the coastal zone of today’s northern North Korea and the RFE, which share several subsistence traits with the Okhotsk culture..

In previous studies, Japanese scholars claimed to have identified a short and a long barley type at Okhotsk and Satsumon culture sites in the Hokkaido Region, which they assigned to the crop’s naked and hulled form, respectively (see [69] and references therein). Based on this differentiation and seed morphology, Yamada and colleagues (e.g. have hypothesised that Okhotsk barley originated from neighbouring regions on the Asian mainland. They found that the highly compact (naked barley) specimens extracted from four Okhotsk culture sites (no. 17–20 in Fig 1CS3 Table) are distinct from the slimmer (hulled) barley (dated to 8–10th century AD) used by Early and Middle Satsumon groups, but similar to grains identified as naked barley found in the early Iron Age to medieval (ca. mid-1st millennium BC–early 13th century AD) sites in southern Primor’e (RFE).
References
Lister DL, Jones H, Oliveira HR, et al. Barley heads east: Genetic analyses reveal routes of spread through diverse Eurasian landscapes. PLoS One. 2018;13(7):e0196652. Published 2018 Jul 18. doi:10.1371/journal.pone.0196652
Leipe C, Sergusheva EA, Mu ̈ller S, Spengler RN, III, Goslar T, Kato H, et al. (2017) Barley (Hordeum vulgare) in the Okhotsk culture (5th–10th century AD) of northern Japan and the role of cultivated plants in hunter–gatherer economies. PLoS ONE 12(3): e0174397. https:// doi.org/10.1371/journal.pone.0174397
PLOS ONE | https://doi.org/10.1371/journal.pone.0174397 March 29, 2017 1 / 27
Further reading about the Okhotsk culture:
Who are the Okhotsk culture people?
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 from mtDNA sequences showed that the Okhotsk people were more closely related to the Nivkhi and Ulchi people among populations of northeastern Asia. In addition, the Okhotsk people had a relatively closer genetic affinity with the Ainu people of Hokkaido, and were likely intermediates of gene flow 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. …
Skeletons of the Okhotsk people share par- ticular morphological characteristics: high and round neu- rocranium, large mandible, flat face, and extremely shallow canine fossa. Kodama (1948) reported that these characters were similar to those of the Aleut people among neigh- boring populations. On the other hand, Suzuki (1958) re- ported that the Okhotsk people were morphologically closer to the Eskimo people than the Aleut. Mitsuhashi and Yamaguchi (1961) found that skeletons of the Okhotsk people have characteristics specific to those of northeastern Asian people. These studies revealed that the morpholog- ical characteristics of the Okhotsk people were clearly different from those of the Ainu people currently living in Hokkaido. In addition, the morphological characteristics of the Okhotsk people are similar to those of the Nivkhi and Ulchi people that are currently distributed around Sakhalin and downstream of the Amur River (Yamaguchi 1974; Ishida 1988, 1996; Kozintsev 1990, 1992). The final con- clusion of the anthropological status of the Okhotsk people, however, has not yet been determined.
On the other hand, evidence of the occurrence of bear-sending ceremonies, as also seen in the Ainu culture, was found from archaeological sites of the Okhotsk culture (Utagawa 2002). In many cases, the skulls of brown bears were enshrined in bone mounds located within houses of the Okhotsk culture. This is a special custom that is not seen in archaeological sites of other cultures in Japan. The ritual is thought to be a proto-type of ‘‘Iomante’’, which was performed as a bear-sending ceremony in the Ainu culture. Ancient DNA analysis of the skulls of brown bears, excavated from an Okhotsk culture site on Rebun island, northern Hokkaido, showed that there were cultural ex- changes through bear cubs between the Okhotsk people and the Epi-jomon people (Masuda et al. 2001), which are archaeologically considered to be direct descendants of the Jomon people, and lived from the third century BC to the seventh century AD in southern Hokkaido. However, the custom of enshrining bears did not occur in the Satsumon culture followed by the Ainu culture. These facts suggest that the Okhotsk culture joined the Satsumon culture in Hokkaido, resulting in establishment of the Ainu culture (Utagawa 2002). …
To investigate the phylogenetic relationships between the Okhotsk people and modern Asian populations, the NJ relationships among the 17 Asian populations were con- structed (Fig. 3). In this phylogenetic tree, the Okhotsk people were clustered with the Nivkhi, Ulchi, Negidal, Koryak, and Even. Among them, the Nivkhi and Ulchi were much closer to the Okhotsk people, and clustered with more than 70% bootstrap values. The close relatedness among the three populations was in congruence with the high degree of sharing of mtDNA haplotypes. On the other hand, the Ainu people were phylogenetically distant from the Okhotsk people (Fig. 3). However, the dA distance
(0.068%, Table 3) between the Okhotsk people and the Ainu was smaller than those between the Okhotsk and other populations except the Nivkhi and Ulchi. Moreover, multidimensional scaling analysis (two-dimensional dis- play, Fig. 4) of the genetic relationships among the 17 Asian populations based on dA distances showed that the Nivkhi, Ulchi, Negidal and Ainu were much closer to the Okhotsk people than the other Asian populations. These findings demonstrate that the Okhotsk people are closely related to modern populations distributed around the Sakhalin and downstream of the Amur River as well as to the Ainu people of Hokkaido.
Among haplotypes 12, 13 and 14 identified from 11 Okhotsk people, a unique combination of four transitional mutations (16189C- 16231C-16266T-16519C for types 12, 13 and 14) was shared and regarded as the motif sequence for mtDNA haplogroup ‘‘Y1’’ reported by Kivisild et al. (2002). Recent studies have shown that most people with haplogroup Y1 are distributed in northern Asia and Siberia (Schurr et al. 1999; Kivisild et al. 2002). Moreover, Adachi et al. (2006) reported that haplogroup Y occurred in the Ainu of Hokkaido but not in the Jomon people of Hokkaido. In the present study, we found that the Okhotsk people shared haplogroup Y1 at a similar frequency (30%, 11/37) to the Nivkhi (35%, 20/57) and the Ulchi (31%, 27/87) (Table 2). The result also suggests that the Okhotsk people are genetically closer to the Nivkhi and Ulchi because of the similarity in the frequencies of haplogroup Y1 between them. Moreover, haplogroup Y1 was shared also by the Ainu (6%, 3/51) (Table 2). That the frequency of Y1 is higher in the Okhotsk people (30%) than the Ainu (6%) suggests gene flow from the Okhotsk people to the Ainu. In addition, other haplotypes clarified to haplogroup Y1 were found: for example, two types from two individuals of the Ulchi, two types from six individuals of the Nivkhi, and one type from seven individuals of the Ainu. When these numbers are included, the same direction of gene flow is still apparent. Tajima et al. (2004) examined mtDNA phylogeny of modern Asian people (not including the Okhotsk people) and reported that there was gene flow from the Nivkhi to the Ainu. The present study demon- strates that the Okhotsk people were an intermediate in the gene flow from the Nivkhi to the Ainu.
In previous morphological studies, Mitsuhashi and Yamaguchi (1961) reported that skeletons of the Okhotsk people had morphological characteristics similar to those of northeastern Asian people. In addition, Yamaguchi(1974) and Ishida (1988, 1996) reported, based on cranial measurements, that the Okhotsk people were closer to populations that are currently distributed downstream of the Amur River. Moreover, Utagawa (2002) reported that evidence for the occurrence of bear-sending ceremonies, as seen in the Ainu culture, was found from archaeological sites of the Okhotsk culture. Rituals using brown bears are thought to be one proto-type of ‘‘Iomante,’’ which has been performed as a bear-sending ceremony in the Ainu culture. Therefore, archaeologists have generally assumed that the Okhotsk culture joined the Satsumon culture (eighth to fourteenth centuries; direct descendants of the Jomon people in Hokkaido) resulting in establishment of the Ainu culture. The direction of gene flow obtained in the present study is in agreement with the interpretation based on previous morphological and archaeological data. These facts demonstrate that the Okhotsk people could have originated from northeastern Asian populations such as the Nivkhi and Ulchi currently living around Sakhalin and downstream of the Amur River, and support the hypothesis that the Okhotsk people could have joined the Satsumon people resulting in the Ainu culture.
Kikuchi (2004) reported that walrus tusks excavated from archaeological sites of the Okhotsk culture could have been brought from the ancient Koryak culture to the Okhotsk culture, because walrus are currently distributed in the Arctic sea and the Bering sea. In the present study, the Okhotsk people were found to have some genetic affinities with the Koryak and the Even living around the Kamchatka peninsula (Table 2; Fig. 3). These facts suggest that there were genetic and cultural exchanges between the Okhotsk people and the Koryak and Even. …
Unique maternal genetic features of the Okhotsk people identified by ancient mtDNA analysis demonstrated their genetic differentiation and some genetic affinity with geographically neighboring modern populations such as the Nivkihi and Ulchi.
Results of this study:
”indicate that the Ainu still retain the matrilineage of the Hokkaido Jomon people. However, the Siberian influence on this population is far greater than previously recog- nized. Moreover, the influence of mainland Japanese is evident, especially in the southwestern part of Hokkaido that is adjacent to Honshu, the main island of Japan.
Discussion: Our results suggest that the Ainu were formed from the Hokkaido Jomon people, but subsequently underwent considerable admixture with adjacent populations. The present study strongly recommends revision of the widely accepted dual-structure model for the population his- tory of the Japanese, in which the Ainu are assumed to be the direct descendants of the Jomon people. …
…, recent morphological and genetic studies (e.g., Hanihara, Yoshida, & Ishida, 2008; Hanihara, 2010; Ishida, Hanihara, Kondo, & Fukumine, 2009; Sato et al., 2009; Shigematsu, Ishida, Goto, & Hanihara, 2004) have indicated that the Siberian influence via the Okhotsk culture people on the Ainu significantly affected the latter’s genetic structure. The Okhotsk culture people are thought to have migrated from north- eastern Eurasia and been distributed in the coastal regions of northern and northeastern Hokkaido as well as southern Sakhalin during the 5th to 13th centuries AD (Amano, 2003). On Hokkaido, this culture was rap- idly diminished by the invasion of Satsumon culture people at the end of the 9th century. As a result, Okhotsk culture had almost disappeared by the beginning of the 10th century. However, in the easternmost part of Hokkaido, the Okhotsk culture transformed into the Tobinitai culture under the strong influence of the Satsumon culture, and this culture con- tinued until the beginning of the Ainu era (Segawa, 2007).
Recently, we also confirmed the considerable genetic influence of the Okhotsk culture people on the formation of the modern-day Ainu.
We found that mitochondrial DNA (mtDNA) haplogroups A, C, and Y, which are shared by the modern-day Siberian populations, Okhotsk cul- ture people, and the modern-day Ainu, are not observed in the Hokkaido Jomon people (Adachi et al., 2011)…
Genetic characteristics of the Ainu mtDNAs
Twenty-one haplogroups and their subhaplogroups were identified in 94 Edo Ainu individuals (Supporting Information Table S1). As described ear- lier, conventionally, the Ainu are considered to be descended from the Hokkaido Jomon people, with little admixture with other populations. Among the haplogroups observed in the Hokkaido Jomon (N9b1, N9b4, N9b*, D4h2, G1b*, M7a2, M7a*; Adachi et al., 2011), haplogroups N9b1, G1b*, and M7a2 are also observed in the Edo Ainu. Above all, haplogroup N9b1, which is the most frequently observed haplogroup in the Hokkaido Jomon people (55.6%, 30 of 54 individuals; Adachi et al., 2011), is also observed at a relatively high frequency (20.2%, 19 of 94 individuals) in
the Edo Ainu. These findings indicate the genetic continuity between the Hokkaido Jomon and the Ainu. This possible genetic continuity is corro- borated by Y chromosome DNA analysis of the modern Ainu. Y chromo- somal DNA haplogroup D1b, which is considered to be a strong candidate for the Jomon paternal lineage, was observed at high frequency in the modern Ainu (Hammer et al., 2006; Tajima et al., 2004).
However, there is a crucial difference between the Hokkaido Jomon people and the Ainu. Four haplogroups (N9b4, N9b*, D4h2, M7a*) are missing in the Edo Ainu, whereas they have 19 haplogroups that are not observed in the Hokkaido Jomon. This raises questions about the conventional hypothesis that the Ainu are the direct descendants of the Hokkaido Jomon people.
To clarify the matrilineal genetic relationship between the Edo Ainu and the other populations, pairwise Fst values between each pair of pop- ulations (Supporting Information Table S2) were calculated from the mtDNA haplogroup frequencies of the Edo Ainu and the 14 ancient and modern-day East Asian and Siberian populations (Table 1, Figure 1). In the frequency-based clustering of compared populations determined by neighbor-joining based on the Fst values described earlier, the Ainu of the Edo era was located almost in the median center between the Hok- kaido Jomon/Udegey cluster and the Lower Amur region cluster includ- ing the Okhotsk culture people (Figure 4). This confirmed our hypothesis (Adachi et al., 2011) that the genetic characteristics of the Ainu are based on the Hokkaido Jomon people and the subsequent input of Lower Amur region Siberian genes through the Okhotsk culture people. However, when examining in detail the haplogroups observed in the Ainu examined in the present study, haplogroups M7a1, M7b1a1a1, D4 (except for D4h2), M8a, Z1a, M9a, F1b, N9a, A5a, and A5c are observed neither in the Hokkaido Jomon nor in the Okhotsk culture people. This suggests the presence of populations other than the Hokkaido Jomon and the Okhotsk culture people that contributed to the formation of the Ainu. Mainland Japanese and the native Siberians are considered to be the major candidates for the origin of these haplogroups because of the proximity of their distributions to Hokkaido. …
the results of mtDNA analysis of the Edo Ainu suggested the multiple origins of this population. This possible multiplicity of origins of the Ainu is considered to be an important factor behind their regional differences.
DISCUSSION
Nowadays, among the theories explaining the population history of the Japanese, the dual-structure model proposed by Hanihara (1991) is the most widely accepted.
However, the results of our study suggest that the later admixture of Ainu with other populations than Jomon people was more consider- able than it was proposed until today.
First, our results showed that the 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.
Moreover, intriguingly, a genetic contribution of the mainland Japanese to the Edo Ainu is evident (28.1%), which is almost as considerable as those of the Jomon and the Okhotsk culture people. As referred to above, 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.
In addition, 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 5 37; Sato et al., 2009), so it is possible that these haplogroups will be identified in the Okhotsk people in further study.
Regional differences of the Ainu
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 the Ainu 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 the individuals in western Hokkaido (this area is included within our category of “southwestern 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 were no regional differences in the frequencies of the Okhotsk-type haplogroups (35.3% in southwestern Hokkaido and 34.9% in northeastern/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.
CONCLUDING REMARKS
In the current study, we clarified that the Ainu were established from the Hokkaido Jomon people and subsequently underwent considerable admixture with adjacent populations. The present study strongly recommends review of the widely accepted dual-structure model regard- ing the population history of the Japanese, in which the Ainu are assumed to be the direct descendants of the Jomon people. However, the causes of the regional differences in the genetic influence of the Okhotsk culture people on the Ainu remain unresolved by mtDNA analysis. Nuclear genome analysis using next-generation sequencing is expected to be helpful to resolve this issue.

Tomb secrets yet to be unlocked from 1,600 years ago

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Click here to see video https://www.bbc.com/reel/embed/p07lq60x

Key-hole shaped tombs in Japan. Video by Matt Dworzańczyk
Image courtesy of Sakai City Government and Sakai City Museum

All kind of megalithic pyramids have awed the ancient local populace for generations. Japan’s ancient Kofun tombs remain relatively unknown or unvisited locations by tourists. This BBC video documentary clip gives a rare English view into the labour-intensive creations of the early centuries of the First Millennium.

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