Japanese wolf

Not to be confused with Hokkaido wolf.
Japanese wolf
Taxidermied specimen, at the National Museum of Nature and Science, Tokyo, Japan
Extinct  (1905)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: C. lupus[1]
Trinomial name
Canis lupus hodophilax
(Temminck, 1839)[2]
Synonyms

The Japanese wolf (Japanese: ニホンオオカミ(日本狼 ー Hepburn: Nihon ōkami) (Canis lupus hodophilax) is an extinct subspecies of the gray wolf that was once endemic to the islands of Honshū, Shikoku, and Kyūshū in the Japanese archipelago. It is also known as the Honshū wolf. Its binomial name derives from the Greek Hodo (path) and phylax (guardian), in reference to Japanese folklore, which portrayed wolves as the protectors of travellers.[4] It was one of two subspecies that were once found in the Japanese archipelago, the other being the Hokkaidō wolf.

Taxonomy and origin

Temminck's depiction of the animal. This may actually represent a yamainu (mountain dog), as the mounted specimen upon which it was based was labeled as such by Siebold, who had encountered both mountain dogs and wolves.[5]

Nomenclature: "ōkami" and "yamainu"

Prior to its classification by Temminck, it had been long recognized in Japan that Honshu was inhabited by two distinct canids; ōkami (wolf) and yamainu (mountain dog), both of which were described by the herbalist Ono Ranzan in his Honzō kōmoku keimō (“An instructional outline of natural studies”) in 1803. He described the ōkami as an edible, but rapacious, greyish-brown animal with a long, ash-colored, white-tipped tail with webbed toes and triangular eyes that would occasionally threaten people if rabid or hungry. In contrast, the yamainu was described as a similar animal, but with speckled yellowish fur, unwebbed toes, a foul odor, and inedible meat.[5]

Ranzan's works were studied by German botanist Philipp Franz von Siebold during his tenure in Dejima. He purchased a female mountain dog and a wolf in 1826, describing both in his notes as distinct, and preparing two sketches illustrating their differences. The skin of the mountain dog was subsequently shipped to the Rijksmuseum van Natuurlijke Historie in the Netherlands and mounted. The specimen, along with Siebold's notes, were used by Dutch zoologist Coenraad Jacob Temminck as references for his scientific classification of the animal in Fauna Japonica (1839). Temminck, however, misinterpreted Siebold's notes distinguishing the wolf and the mountain dog and treated the two as synonyms. In 1842, he wrote a longer description, still confounding the two names, and producing a sketch a "wolf" based on Siebold's mounted mountain dog specimen.[5]

Skeletal and genetic findings

Mounted skeleton, National Museum of Nature and Science

The Japanese wolf,[6][7][8][9]:42 or Honshū wolf,[8] (Canis lupus hodophilax Temminck, 1893)[1][3] is a subspecies of the gray wolf (Canis lupus). Skeletal remains of the Japanese wolf have been found in archaeological sites dating from the Jomon Period (10,000 to 250 B.C).[10][11]

The Japanese wolf was not the world's smallest wolf. The cranial length of the adult Arab wolf (Canis lupus arabs) measures on average 200.8 mm, which is smaller than most wolves. Specimens of the Japanese wolf were measured between 193.1 mm and 235.9 mm and it was uncertain if these were all from adults.[9]:53 In the mandible, M1 (molar tooth) is relatively larger than in any other canid species.[12][11] An examination in 1991 found one specimen's condylobasal length (a measure of skull length) to be 205.2mm, and the Alveolar length of P4 (the fourth maxillary premolar or carnassial tooth) to be 20.0mm (left) and 21.0mm (right).[13] In 2009, an osteological study declared that the skull of the Japanese wolf was between 206.4 mm to 226.0 mm in total length, and that morphological characters alone were not sufficient to distinguish the Japanese wolf from large domesticated dogs, such as the Akita breed.[11] Remains of the wild native canine dating from the late Edo period (1603 and 1868), the Yama-Inu, has occasionally been confused with the Japanese wolf because of the osteological similarities between the two.[11][9]:40[14]

The Japanese wolf inhabited Kyushu, Shikoku, and Honshu islands[11][9]:42 but not Hokkaido Island.[11] This indicates that its ancestor may have migrated from the Asian continent through the Korean Peninsula into Japan.[11][8] From mitochondrial DNA analysis, the phylogenetic tree reveals a long branch separating the Japanese wolf from other gray wolf populations, suggesting a long period of isolation in the Japanese archipelago. The wolf appears to have arrived in Japan during the Late Pleistocene between 25,000–125,000 years ago. The analysis indicated that the Japanese wolf belongs to the ancient mDNA haplogroup 2[8] (solely represented today by the Italian wolf and scattered pockets of other wolves across Eurasia),[15] while the Hokkaido wolf belongs to mDNA haplogroup 1, which suggests that the Japanese wolf was the first arrival on the Japanese archipelago with the Hokkaido wolf arriving more recently from the north.[8]

Specimen in Ueno Zoo
See further Gray wolf haplogroups

An examination of sequences from 113 ancient Canis specimens from China and Russia did not match, which indicated that none of these specimens were the ancestors of the Japanese wolf.[16]

Analyses of the mitochondrial DNA of 1576 dogs worldwide revealed that one Kishu[11][8] and one Siberian husky[11][8] possessed the same haplotype as a Japanese wolf, indicating past cross-breeding.[11][17][18] A more-refined study of Japanese wolf mitochondrial DNA showed that they could be further divided into two separate groups, and that the sequences from one Kishu, one Siberian husky and one Shiba Inu could also be divided into the two groups.[16] These dogs correspond to clade F of the mDNA phylogenetic tree among worldwide dogs,[19][20][21] with clade F haplogroup dogs originating from a rare admixture between male dogs and more than one female ancestor of Japanese wolves, which have contributed to the dog gene pool.[8][19][20]

See further Dog-Wolf hybridization

Range

Carl Hubert de Villeneuve's sketch of Siebold's captive specimen, probably the only authentic European depiction of the species unambiguously identified as a wolf rather than a "mountain dog"[5]

The Japanese wolf inhabited Kyushu, Shikoku, and Honshu islands[11][9]:42 but not Hokkaido Island.[11] The remains of a 28,000 year old wolf specimen from the Yana River on the northern coast of arctic Siberia matched the mDNA haplotype of the Japanese wolf, which indicates that they shared common ancestry[7] and a wider distribution.

Physical characteristics

Illustration from 1881 labelled "Canis hodophylax or Japanese wolf",[22] also likely to be a yamainu (mountain dog) rather than the Japanese wolf

Canis lupus hodophilax was described by Temminck in 1839 as smaller than Canis lupus lupus (Linnaeus 1758) and of shorter legs, with its coat smooth and short.[23] The Japanese wolf was smaller in size compared to the Hokkaido wolf and other gray wolves from the Asian and North American continents.[24]

There are four mounted specimens believed to be Canis lupus hodophilax located at: the National Museum of Nature and Science, Japan; University of Tokyo, Japan; Wakayama University, Japan; Siebold Collection, and the National Museum of Natural History, Leiden, Netherlands.[11]

History

The Japanese wolf is considered to be extinct[25] as the last specimens were recorded at Higashi-Yoshino village in Nara Prefecture, Japan in 1905.[11]

The Japanese wolf population began to decrease in 1732 when rabies, first reported in Kyūshū and Shikoku, was introduced to the area they inhabited. It affected different wolf populations all through the nineteenth century. It is believed that local domestic dogs in the regions may have transported the disease. Along with intense human persecution, the wolves proceeded into extinction.[26][27] Some interpretations of the Japanese wolf's extinction stress the change in local perceptions of the animal: rabies-induced aggression and deforestation of the wolf's habitat forced them into conflict with humans, and this led to them being targeted by farmers.[4] Other sources say the wolves were killed off as a national policy.[28]

Sightings of "short-legged dog like beasts", proposed to be the Japanese wolf, have been claimed since the time of its extinction until the last claim in 1997, but none of these have been verified. A claim in 2000 was dismissed as a hoax. Some Japanese zoologists believe that these reports "merely derive from misidentification of feral dogs".[29]

Culture

Kitayama moon by Yoshitoshi.[30] The wolf drawings were taken from a book illustrated by Kuniyoshi, who was Yoshitoshi’s teacher.[31]
Statue in Japan, with the inscribed haiku: I walk/ With that wolf/ That is no more

The Japanese wolf is a prominent figure in Japanese folklore and culture; in ancient Japan, the wolf was seen through Shintoism as a kami (god) inhabiting a sacred and chaotic landscape which contrasted with the rigid etiquette of court life.[9]:66-69 The mountains of Japan, seen as a dangerous, deadly place, were highly associated with the wolf, which was believed to be their protector and guardian. Many mountain villages, such as Okami’iwa (Wolf Rock) and Okamitaira (Wolf Plateau), are named after the wolf; this could be due to a sighting at the location, or a simple homage to the species.[4] Sightings of the Japanese wolf were very rare, and the wolf was described as being more of a spirit entity protecting travelers. Some legends bring this to a whole new level, where abandoned infants are found and raised by the wolves. The wolves also protect surrounding villages from the dangerous wildlife.[26] The spirit form of the Japanese wolf, which has been worshiped since ancient times, is called makami (真神). It is believed to understand human speech, reward good, and punish evil. Makami shrines are found in Saitama, Shizuoka, and Tokyo.

The wolf was afforded a benign place in Japanese folklore and religious traditions: the clan leader Fujiwara no Hidehira was said to have been raised by wolves, and the wolf is often symbolically linked with mountain kami in Shinto. The most famous example is the wolf kami of Mitsumine Shrine in the town of Chichibu in Saitama Prefecture.

Some villages have wolf charms called shishiyoke, that protected their village and their crops against wild boar. In addition, other protective shrines can be found in many villages, especially on the Kii Peninsula. In some villages, such as in Gifu prefecture, the skull of the wolf was used as the charm for both protection as well as curing possessed villagers. In addition to protecting the crops, the wolf may leave prey for villagers.[4] A tradition called inu no ubumimai consists of giving a mother Japanese wolf rice when she gives birth to a cub. In return, the wolf would protect the village and assist in danger, or leave the village in times of famine.[26]

Some legends portray the Japanese wolf as being prophetic creatures. In the Tamaki Mountains the location of a tree called “the cypress of dog-howls” is said to be the site where wolves howled before a flood in 1889, warning the villagers.[4] The Japanese wolf was not commonly killed by villagers, and attacks were rare. Some folklore states that the killer of a Japanese wolf faced punishment from the spiritual world. Legends also describes the wolf as being concealed by the environment, and that its fur changes with the seasons to further camouflage itself. In many local cultures, therefore, the Japanese wolf is believed not to be extinct; it was always difficult to find.[26]

References

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  1. 1 2 Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ↑ Temminck, C. J. (1839) Over de Kennis en de Verbreiding der Zoogdieren van Japan. Tijidschrift voor Natuurlijke Geschiedenis en Physiologie, pt5, 274-293 - refer page 284
  3. 1 2 3 Smithsonian - Animal Species of the World database. "Canis lupus hodophilax".
  4. 1 2 3 4 5 Knight, John (1997). "On the Extinction of the Japanese Wolf". Asian Folklore Studies (Nanzan University) 56 (1): 129–159. doi:10.2307/1178791. Retrieved January 24, 2014. Stable URL: http://www.jstor.org/stable/1178791
  5. 1 2 3 4 Funk, H. (2015). "A re-examination of C. J. Temminck's sources for his descriptions of the extinct Japanese wolf". Archives of natural history 42 (1): 51–65. ISSN 0260-9541.
  6. ↑ "Canis lupus hodophilax (Japanese wolf)". NCBI.NLM.NIH.gov. National Center for Biotechnology Information, U.S. National Institutes of Health.
  7. 1 2 Lee, E. (2015). "Ancient DNA analysis of the oldest canid species from the Siberian Arctic and genetic contribution to the domestic dog". PLoS ONE 10 (5): e0125759. doi:10.1371/journal.pone.0125759. PMC 4446326. PMID 26018528.
  8. 1 2 3 4 5 6 7 8 Matsumura, Shuichi; Inoshima, Yasuo; Ishiguro, Naotaka (2014). "Reconstructing the colonization history of lost wolf lineages by the analysis of the mitochondrial genome". Molecular Phylogenetics and Evolution 80: 105–12. doi:10.1016/j.ympev.2014.08.004. PMID 25132126.
  9. 1 2 3 4 5 6 Walker, Brett (2008). The Lost Wolves of Japan. University of Washington Press.
  10. ↑ Shigehara, N; Hongo, H (2000). "Dog and wolf remains of the earliest Jomon period at Torihama site in Fukui Prefecture" (in Japanese) 2. Torihama-Kaizuka-Kennkyu: 23–40.
  11. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Ishiguro, Naotaka; Inoshima, Yasuo; Shigehara, Nobuo (2009). "Mitochondrial DNA Analysis of the Japanese Wolf (Canis Lupus Hodophilax Temminck, 1839) and Comparison with Representative Wolf and Domestic Dog Haplotypes". Zoological Science 26 (11): 765. doi:10.2108/zsj.26.765. PMID 19877836.
  12. ↑ Miyamoto F, Maki I (1983) On the repaired specimen of Japanese wolf (Canis lupus hodophilax Temminck) and its skull newly taken out. Bull Fac Ed Wakayama Univ Nat Sci 32: 9–16 (in Japanese with English abstract)
  13. ↑ Miyamoto, F. (1991). On the skull of Japanese wolf (Canis hodophilax Temminck) taken out from the mounted specimen preserved in Wakayama University. Bull. Fac. Ed. Wakayama Univ. Nat. Sci. 39: 55–60. (in Japanese with English abstract)
  14. ↑ Obara I, Nakamura K (1992) Notes on a skull of so-called “Yama-Inu” or wild canine preserved in the Minamiashigara municipal folklore museum. Bull Kanagawa Pref Mus Nat Sci 21: 105–110 (in Japanese with English abstract)
  15. ↑ Pilot, M. G.; Branicki, W.; Jędrzejewski, W. O.; Goszczyński, J.; Jędrzejewska, B. A.; Dykyy, I.; Shkvyrya, M.; Tsingarska, E. (2010). "Phylogeographic history of grey wolves in Europe". BMC Evolutionary Biology 10: 104. doi:10.1186/1471-2148-10-104. PMC 2873414. PMID 20409299.
  16. 1 2 Ishiguro, Naotaka; Inoshima, Yasuo; Yanai, Tokuma; Sasaki, Motoki; Matsui, Akira; Kikuchi, Hiroki; Maruyama, Masashi; Hongo, Hitomi; Vostretsov, Yuri E.; Gasilin, Viatcheslav; Kosintsev, Pavel A.; Quanjia, Chen; Chunxue, Wang (2016). "Japanese Wolves are Genetically Divided into Two Groups Based on an 8-Nucleotide Insertion/Deletion within the mtDNA Control Region". Zoological Science 33 (1): 44–9. doi:10.2108/zs150110. PMID 26853868.
  17. ↑ Okumura N, Ishiguro N, Nakano M, Matsui A, Sahara M (1996) Intra- and interbreed genetic variations of mitochondrial DNA major noncoding regions in Japanese native dog breeds (Canis familiaris). Anim Genet 27: 397–405
  18. ↑ Takahashi S, Miyahara K, Ishikawa H, Ishiguro N, Suzuki M (2002) Lineage classification of canine inheritable disorders using mitochondrial DNA haplotypes. J Vet Med Sci 64: 255–259
  19. 1 2 Pang, J. (2009). "mtDNA data indicate a single origin for dogs south of Yangtze River, less than 16,300 years ago, from numerous wolves". Molecular Biology and Evolution 26 (12): 2849–64. doi:10.1093/molbev/msp195. PMC 2775109. PMID 19723671.
  20. 1 2 Duleba, Anna; Skonieczna, Katarzyna; Bogdanowicz, Wiesław; Malyarchuk, Boris; Grzybowski, Tomasz (2015). "Complete mitochondrial genome database and standardized classification system for Canis lupus familiaris". Forensic Science International: Genetics 19: 123–129. doi:10.1016/j.fsigen.2015.06.014.
  21. ↑ Savolainen, P. (2002). "Genetic evidence for an East Asian origin of domestic dogs". Science 298 (5598): 1610–3. doi:10.1126/science.1073906. PMID 12446907.
  22. ↑ Brauns, D., 1881. Canis Hodophylax, or Japanese wolf. Chrysanthemum, A Monthly Magazine for Japan and the Far East, 1 (January to December, 1881): 66-67, plate before page 66. Kelly & Co., Yokohama.
  23. ↑ Mech, L David (1970) "The wolf: the Ecology and Behavior of an Endangered Species", published for the American Museum of Natural History by the Natural History Press, pages 353
  24. ↑ Ishiguro, Naotaka; Inoshima, Yasuo; Shigehara, Nobuo; Ichikawa, Hideo; Kato, Masaru (2010). "Osteological and Genetic Analysis of the Extinct Ezo Wolf (Canis Lupus Hattai) from Hokkaido Island, Japan". Zoological Science 27 (4): 320. doi:10.2108/zsj.27.320. PMID 20377350.
  25. ↑ Claudio Sillero-Zubiri (2009). "Family Canidae (Dogs)". In Don E. Wilson, Russell A. Mittermeier. Handbook of the Mammals of the World - Volume 1:Carnivores. Lynx Edicions in association with Conservation International and IUCN. p. 413.
  26. 1 2 3 4 "Lost Wolves of Japan". SciLogs. Retrieved 23 October 2012.
  27. ↑ "The Honshu Wolf". Bib.ge. Retrieved 23 October 2012.
  28. ↑ Chester Dawson (December 28, 2012). "If the Japanese Diet Included Deer, It Might Keep Wolves From the Door". Wall Street Journal.
  29. ↑ Shuker, Karl P. N. (2003). The beasts that hide from man: seeking the world's last undiscovered animals. New York: Paraview Press. pp. 218–221. ISBN 1-931044-64-3.
  30. ↑ Kitayama moon - Toyohara Sumiaki, from the series One hundred aspects of the moon by Tsukioka Yoshitori. Art Gallery of New South Wales, Australia
  31. ↑ Japan Print Gallery, Notting Hill Gate, London
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