|Taxidermied specimen, at the National Museum of Nature and Science, Tokyo, Japan|
|Canis lupus hodophilax
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. It was one of two subspecies that were once found in the Japanese archipelago, the other being the Hokkaidō wolf.
Taxonomy and origin
Nomenclature: "ōkami" and "yamainu"
Before Temminck classified it, 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.
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 of a "wolf" based on Siebold's mounted mountain dog specimen.
Skeletal and genetic findings
The Japanese wolf, or Honshū wolf, (Canis lupus hodophilax Temminck, 1893) is a subspecies of the gray wolf (Canis lupus). Skeletal remains of the Japanese wolf have been found in archaeological sites, such as Torihama shell mound, dating from the Jōmon period (10,000 to 250 B.C).
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. In the mandible, M1 (molar tooth) is relatively larger than in any other canid species. 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). 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. 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.
The Japanese wolf inhabited Kyushu, Shikoku, and Honshu islands but not Hokkaido Island. This indicates that its ancestor may have migrated from the Asian continent through the Korean Peninsula into Japan. The phylogenetic tree generated from its mitochondrial DNA sequences revealed a long branch that separated the Japanese wolf from other gray wolf populations and that it belongs to the ancient mDNA haplogroup 2 (represented today by the Italian wolf and scattered pockets of other wolves across Eurasia), while the Hokkaido wolf belongs to mDNA haplogroup 1 and this suggests that the Japanese wolf was the first arrival on the Japanese archipelago with the Hokkaido wolf arriving more recently from the north. The wolf was estimated to have arrived in Japan during the Late Pleistocene between 25,000–125,000 years ago, however a more recent study that looked at the past sea levels of the Korean Strait together with the timing of the Japanese wolf sequences indicated that it arrived to the southern islands less than 20,000 YBP.
- See further: Evolution of the wolf - North America and Japan
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.
Analyses of the mitochondrial DNA of 1576 dogs worldwide revealed that one Kishu and one Siberian husky possessed the same haplotype as a Japanese wolf, indicating past cross-breeding. 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. These dogs correspond to clade F of the mDNA phylogenetic tree among worldwide dogs, 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.
- See further Dog-Wolf hybridization
The Japanese wolf inhabited Kyushu, Shikoku, and Honshu islands but not Hokkaido Island. 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 and a wider distribution.
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. The Japanese wolf was smaller in size compared to the Hokkaido wolf and other gray wolves from the Asian and North American continents. It stood 56–58 cm at the withers.
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.
The Japanese wolf is considered to be extinct as the last specimens were recorded at Higashi-Yoshino village in Nara Prefecture, Japan in 1905. 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".
In 713, the wolf first appeared on record in Kofudoki itsubun (Lost writings on ancient customs). From 967, historical records indicated the wolf's preference for preying on horse, either wild horses or those in pastures, stables, and villages. In 1701, a lord introduced the first wolf bounty and by 1742 the first professional wolf hunters were using firearms and poison. In 1736, rabies appeared among dogs in eastern Japan, indicating that it had entered from China or Korea, then spread across the nation. Shortly after it spread to the wolf population, turning some wolves from simple horse predators to man-killers that lead to organized wolf hunts. Killing wolves became a national policy under the Meiji Restoration, and within one generation the Japanese wolf was extinct.
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.
In the Shinto belief, the ōkami ("wolf") is regarded as a messenger of the kami spirits and also offers protection against crop raiders such as the wild boar and deer. Wild animals were associated with the mountain spirit Yama-no-kami. 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 Okamiiwa ("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.
There are an estimated 20 Shinto wolf shrines on Honshu alone. The most famous national shrine is located at Mitsumine in Chichibu, Saitama Prefecture and there are a number of smaller wolf shrines on the Kii Peninsula, including the Tamaki Shrine and the Katakati Shrine at Totsukawa village.
In Japanese folklore, there is the widely recorded belief of the okuriōkami ("escort wolf") that followed someone walking alone through a forest at night until they reach their home without doing them any harm. An offering was sometimes made for this escort. Another belief was of wolves that raised an infant who had been abandoned in the forests of the Kii Peninsula, and later became the clan leader Fujiwara no Hidehira. Another belief from the Kanto area of eastern Japan was that feeding an infant wolf's milk would make them grow up strong. 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 immediately before a flood in 1889 warning the villagers, and before the great earthquake of 1923 even though the wolf was extinct by that time. Another belief was the "wolf notification" where a traveller does not return home, then a wolf comes to their home and makes a sad howling that signalled their death.
Some villages had wolf charms called shishiyoke that were believed to protect their village and their crops against wild boar. Wolf fangs, hide, and hair were carried by travelers to ward of evil spirits, and wolf skulls were kept in some home shrines to ward off misfortune. 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.
- 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.
- 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
- Smithsonian - Animal Species of the World database. "Canis lupus hodophilax".
- 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
- Kokugo Dai Jiten, Revised Edition 1988 (in Japanese), Tōkyō:Shogakukan
- 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. doi:10.3366/anh.2015.0278.
- "Canis lupus hodophilax (Japanese wolf)". NCBI.NLM.NIH.gov. National Center for Biotechnology Information, U.S. National Institutes of Health.
- 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. PMC . PMID 26018528. doi:10.1371/journal.pone.0125759.
- 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. PMID 25132126. doi:10.1016/j.ympev.2014.08.004.
- Walker 2008, p. 42.
- 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.
- 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): 765–70. PMID 19877836. doi:10.2108/zsj.26.765.
- Walker 2008, p. 53.
- 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)
- 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) 
- Walker 2008, p. 40.
- 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) 
- Koblmüller, Stephan; Vilà, Carles; Lorente-Galdos, Belen; Dabad, Marc; Ramirez, Oscar; Marques-Bonet, Tomas; Wayne, Robert K.; Leonard, Jennifer A. (2016). "Whole mitochondrial genomes illuminate ancient intercontinental dispersals of grey wolves (Canis lupus)". Journal of Biogeography. doi:10.1111/jbi.12765.
- 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. PMC . PMID 20409299. doi:10.1186/1471-2148-10-104.
- 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. PMID 26853868. doi:10.2108/zs150110.
- 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.
- 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.
- 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. PMC . PMID 19723671. doi:10.1093/molbev/msp195.
- 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.
- Savolainen, P. (2002). "Genetic evidence for an East Asian origin of domestic dogs". Science. 298 (5598): 1610–3. PMID 12446907. doi:10.1126/science.1073906.
- 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. 
- 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
- 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): 320–4. PMID 20377350. doi:10.2108/zsj.27.320.
- Ishiguro, Naotaka (2011). "Phylogenetic analysis of extinct wolves in japan" (PDF). Gifu University, Japan. p. 11. in Japanese, measurements in English
- 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.
- 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.
- Walker 2008, p. 101-106.
- Walker 2008, p. 113-119.
- Walker 2008, p. 128.
- Kitayama moon - Toyohara Sumiaki, from the series One hundred aspects of the moon by Tsukioka Yoshitori. Art Gallery of New South Wales, Australia 
- Japan Print Gallery, Notting Hill Gate, London
- Walker 2008, p. 7.
- Knight 2006, p. 4197-8.
- Knight 2006, p. 205.
- Knight 2006, p. 206.
|Wikispecies has information related to: Canis lupus hodophilax|
|Wikimedia Commons has media related to Canis lupus hodophilax.|