Japanese wolf

The Japanese wolf (Japanese: ニホンオオカミ(日本狼, 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 Hodos (path) and phylax (guardian), in reference to Japanese folklore, which portrayed wolves as the protectors of travellers.[3] It was one of two subspecies that were once found in the Japanese archipelago, the other being the Hokkaido wolf.

Not to be confused with Hokkaido wolf.

Japanese wolf
Mounted 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
Subspecies:
†C. l. hodophilax
Trinomial name
Canis lupus hodophilax
(Temminck, 1839)[1]
Synonyms

Etymology

The name ōkami (wolf) is derived from the Old Japanese öpö-kamï, meaning "great-spirit".[4] In the Shinto religion, wild animals were associated with the mountain spirit Yama-no-kami.[3]

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

Nomenclature: "ōkami" and "yamainu"

Before Dutch zoologist Coenraad Jacob 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.[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 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.[5]

Skeletal and genetic findings
Mounted skeleton, National Museum of Nature and Science
Skull in the Ueno Zoo, Japan
Mounted specimen in Ueno Zoo

The Japanese wolf,[6][7][8][9] or Honshū wolf,[8] (Canis lupus hodophilax Temminck, 1893)[10][2] 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 mounds, dating from the Jōmon period (10,000 to 250 B.C).[11][12]

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.[13] In the mandible, M1 (molar tooth) is relatively larger than in any other canid species.[12][14] 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).[15] 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.[12] 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.[16][12][17]

The Japanese wolf inhabited Kyushu, Shikoku, and Honshu Islands[9][12] but not Hokkaido Island.[12] This indicates that its ancestor may have migrated from the Asian continent through the Korean Peninsula into Japan.[8][12][18] 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[8][18] (represented today by the Italian wolf and scattered pockets of other wolves across Eurasia[19]), 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,[8] 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.[18]

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.[20]

Analyses of the mitochondrial DNA of 1576 dogs worldwide revealed that one Kishu[8][12] and one Siberian husky[8][12] possessed the same haplotype as a Japanese wolf, indicating past cross-breeding.[12][21][22] 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.[20] These dogs correspond to clade F of the mDNA phylogenetic tree among worldwide dogs,[23][24][25] 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][23][24]

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[9][12] but not Hokkaido Island.[12] 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",[26] 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.[27] The Japanese wolf was smaller in size compared to the Hokkaido wolf and other gray wolves from the Asian and North American continents.[28] It stood 56–58 cm at the withers.[29]

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.[12]

History
Full moon in Mushasi by Yoshitoshi 1890

The Japanese wolf is considered to be extinct[30] as the last Japanese wolf was captured and killed at Washikaguchi of Higashiyoshino village in Honshu Nara Prefecture, Japan on January 23, 1905.[12] 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".[31]

In AD 713, the wolf first appeared on record in Kofudoki itsubun (Lost writings on ancient customs). From AD 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.[32] 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 led to organized wolf hunts.[33] Killing wolves became a national policy under the Meiji Restoration, and within one generation the Japanese wolf was extinct.[34]

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.[3]

Culture
Kitayama Moon by Yoshitoshi.[35] The wolf drawings were taken from a book illustrated by Yoshitoshi's teacher, Kuniyoshi.[36]
The Japanese wolf memorial stands by the banks of the Takami River in Higashiyoshino, Nara and is a replica of the last wolf killed by hunters nearby in 1905.[37] Inscribed is the haiku: I walk/ With that wolf/ That is no more
Japanese wolf mounted in Ueno Zoo, Japan (Wakayama University possession)

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.[3]

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.[38]

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.[39] 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,[3] and before the great earthquake of 1923 even though the wolf was extinct by that time.[40] 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.[40]

Some villages had wolf charms called shishiyoke that were believed to protect their village and their crops against wild boar.[3] 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.[40] 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.[3]

Claimed post-extinction records

Despite the status, there have been various reports of canines resembling Canis lupus hodophilax throughout the 20th century and in the 21st century including a case by foreign tourists.[41] Three of these, a kill within Fukui Castle in 1910[42][43] and two sightings from Chichibu in 1996[41][44] and nearby Mount Sobo in 2000,[45][46] involved closely taken images of each animals and scientific investigations. These cases triggered debates both for and against the identities of the animals; however, affirmative biologists claimed morphological correspondences of all to Canis lupus hodophilax rather than misidentifications of feral animals such as a Eurasian wolf for the 1910 capture or Shikoku dog for the sighting in 2000. For 1910 record, scientists agreed that this was a Canis while some pointed the possibility of a Eurasian wolf that fled from a mobile zoo four or five days before; however, a staff of the zoo checked the corpse and confirmed that the animal captured was different.[42]

gollark: Metatrolling can never end well.
gollark: Hmm, they need moar views.
gollark: _prepares eggs for sending off_
gollark: I have 5 left which I'm saving for some nebulous "later".
gollark: _perpetually needs more reds_

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Further reading
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