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Mamushi togurosugata

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Subordo: Serpentes
Infraordo: Caenophidia
Superfamilia: Viperoidea

Familia: Viperidae
Subfamilia: Crotalinae
Genus: Gloydius
Species: Gloydius blomhoffii

Gloydius blomhoffii (Boie, 1826)

Holotype: ZMA RENA.15179, collected by J.C. Blomhoff between 1817–1824.
Type locality: “Japan”.


Trigonocephalus blomhoffii Boie, 1826: 214 [original combination]
Halys blomhoffii — Günther, 1864: 393 [subsequent combination]
Ancistrodon blomhoffii — Boulenger, 1890: 140 [subsequent combination]
Agkistrodon blomhoffii — Stejneger, 1907: 457 [subsequent combination]
Gloydius blomhoffii — Hoge & Romano-Hoge, 1980: 195 [subsequent combination]


Trigonocephalus affinis Gray, 1849: 14 [status unclear]
Holotype: BMNH 1946.1.20.1 (formerly BMNH i.7.2a), ♂, presented by Capt. Edward Belcher.
Type locality: unknown, possibly from “Ishigaki shima, in the Yaeyama group” fide Stejneger (1907: 461).

Trigonocephalus blomhoffii megaspilus Cope, 1860: 336
Holotype: ANSP 136, collected in Perry Expedition between 1852–1854.
Type locality: unknown, probably Japan.

Native distribution areas

Japan: Hokkaido, Honshu, Shikoku, Kyushu and the satellite islands.
Russia: Kunasir Island.

Primary references

Boie, H. 1826. Merkmale einiger japanischen Lurche. Isis von Oken 18(2): 203–216. BHL Reference page.
Gray, J.E. 1849. Catalogue of the specimens of snakes in the collection of the British Museum. Edward Newman: London. i–xv + 1–125. BHL Reference page.
Cope, E.D. [1859] 1860. Catalogue of the venomous serpents in the Museum of the Academy of Natural Sciences of Philadelphia, with notes on the families, genera and species. Proceedings of the Academy of Natural Sciences of Philadelphia 11: 332–347. BHL Reference page.
Stejneger, L.H. 1907. Herpetology of Japan and adjacent territory. Bulletin of the United States National Museum 58: xx + 1–577, 35 pls. BHL Reference page.

Additional references

Orlov, N.L., Sundukov, Y. & Kropachev, I. 2014. Distribution of Pitvipers of «Gloydius blomhoffii» Complex in Russia with the First Records of Gloydius blomhoffii blomhoffii at Kunashir Island (Kuril Archipelago, Russian Far East). Russian Journal of Herpetology 21(3): 169–178. DOI: 10.30906/1026-2296-2014-21-3-169-178 [nonfunctional] Broken access Reference page.


Uetz, P. & Hallermann, J. 2021. Gloydius blomhoffii. The Reptile Database. Accessed on 6 May 2019.
Kidera, N. & Ota, H. 2018. IUCN: Gloydius blomhoffii (Least Concern). The IUCN Red List of Threatened Species 2018: e.T192065A2035458. DOI: 10.2305/IUCN.UK.2018-2.RLTS.T192065A2035458.en

Vernacular names
English: Mamushi
日本語: ニホンマムシ
한국어: 살무사

Gloydius blomhoffii, commonly known as the mamushi,[3] Japanese moccasin, Japanese pit viper, Qichun snake, Salmusa or Japanese mamushi,[4] is a venomous pit viper species found in Japan. It was once considered to have 4 subspecies, but it is now considered monotypic.[5]

This species, the yamakagashi (Rhabdophis tigrinus) and the Okinawan habu (Protobothrops flavoviridis) are the most venomous snakes in Japan.[6] Every year, 2000–3000 people in Japan are bitten by a mamushi. Bitten victims typically require one week of treatment in a hospital. Severe bites require intensive care, and approximately 10 victims die annually.[7][8]


The specific name, blomhoffii, is in honor of Jan Cock Blomhoff, who was director of the Dutch trading colony in Nagasaki, Japan from 1817 to 1824.[9]

The average length of mature individuals is 45–81 cm (17¾-31⅞ inches); the longest specimen ever recorded had a length of 91 cm (36 in).[3]

The body pattern consists of a pale gray, reddish-brown, or yellow-brown background, overlaid with a series of irregularly-shaped lateral blotches. These blotches are bordered with black and often have lighter centers. The head is dark brown or black, with beige or pale-gray sides.[3]
Sign warning for mamushi in Kyoto, Japan
Common names

The common name in English is mamushi,[3] or Japanese mamushi.[4] The common name in Japanese is mamushi (蝮). In Korea, it is known as Korean: 살무사; RR: salmusa or Korean: 살모사; RR: salmosa. In China, it is known as the Qichun snake (七寸子) or soil snake/viper (土巴蛇、土蝮蛇、土夫蛇、土公蛇).
Geographic range

It is found in Japan. According to Gloyd and Conant,[10] there is no evidence to support claims that this species occurs in the Ryukyu Islands. The type locality given is "Japan".[2]

It occurs in a range of habitats, including swamps, marshes, meadows, open woodland, rocky hillsides, and montane rock outcroppings.[3]
A mamushi lurking in a bush a little above ground-level, waiting to ambush passing prey

It is typically an ambush predator that uses its excellent camouflage to hide itself in vegetation or leaf litter. It hunts and eats mainly rodents, but also small birds, lizards, and insects. It is often found in and around farmland due to the associated rodent populations.[3]

The venom of this species varies very little among Japan in terms of both its potency and its effects.[11] According to Yoshimitsu (2005), this species and the Okinawan habu (Protobothrops flavoviridis), another pit viper, are the most venomous snakes in Japan.[6] The venom's lethality as measured by LD50 in mice following intraperitoneal injection is in the range 0.3 mg/kg[12] to 1.22 mg/kg.[13] The venom mostly contains haemolytic toxins, but it also has two neurotoxins—an alpha-toxin that is a post-synaptic inhibitor and a beta-toxin that is a pre-synaptic inhibitor.[13] Because the beta-toxin acts pre-synaptically, its effects cannot be blocked or treated by anticholinesterases.[13] The venom contains an anticoagulant, mamushi L-amino-acid oxidase (M-LAO).[14] It also contains the peptide ablomin which is highly similar in amino acid sequence to that of the venom, helothermine, of the Mexican beaded lizard (Heloderma horridum).[15]
Treatments for envenomations

There is an effective antivenom manufactured in both Japan and China.[11] Its effectiveness is increased when co-administered with a serine protease inhibitor such as FOY (see, e.g. Camostat).[16] In common with many other venomous snakes, the mamushi is highly resistant to its own venom because of various neutralising factors present in its sera including phospholipase A2 (PLA2) inhibitors; these and other inhibitors are the target of antivenom development.[17]

Every year, 2000-3000 people in Japan are bitten by mamushi, severe bites require intensive care, and approximately 10 victims die.[7] There have been case reports of kidney failure,[18] visual disturbances,[19] palsy, and miscarriage in pregnant women.[20]

In one study in Japan, mamushi bite victims required a median duration of 7 days of hospital treatment followed by a median of 31 days of out-patient treatment; the time to achieve a full recovery was even longer, taking up to several months.[8] The treatment protocol involved incision of the wound for exclusion of the venom, and injection of mamushi antivenom.[8]

This species is similar to the cottonmouths and copperheads (Agkistrodon sp.) of the Americas, and it was long considered part of the same group (see synonymy).[2]

Kidera, N. & Ota, H. 2018. Gloydius blomhoffii. The IUCN Red List of Threatened Species 2018: e.T192065A2035458. Downloaded on 06 June 2021.
McDiarmid RW, Campbell JA, Touré T (1999). Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Washington, District of Columbia: Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).
Mehrtens JM (1987). Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. ISBN 0-8069-6460-X.
Gumprecht A, Tillack F, Orlov NL, Captain A, Ryabov S (2004). Asian Pitvipers. First Edition. Berlin: Geitje Books. 368 pp. ISBN 3-937975-00-4.
"Gloydius blomhoffii ". Integrated Taxonomic Information System. Retrieved 19 May 2007.
Yoshimitsu, M (2005). "Animal and Snake Bites". Japanese Journal of Pediatric Surgery (in Japanese). 37 (2): 207–15. ISSN 0385-6313.
Okamoto, Osamu; Oishi, Masaki; Hatano, Yutaka; Kai, Yoshitaka; Goto, Mizuki; Kato, Aiko; Shimizu, Fumiaki; Katagiri, Kazumoto; Fujiwara, Sakuhei (2009). "Severity factors of Mamushi (Agkistrodon blomhoffii) bite". The Journal of Dermatology. 36 (5): 277–83. doi:10.1111/j.1346-8138.2009.00638.x. PMID 19382998. S2CID 33668614.
Shigeta, Masatoshi; Kuga, Takayuki; Kudo, Junichi; Yamashita, Akimasa; Fujii, Yasuhiro (2007). "Clinical Study of Mamushi Viper Bites in 35 Cases". Journal of the Japanese Association of Rural Medicine. 56 (2): 61–7. doi:10.2185/jjrm.56.61.
Beolens, Bo; Watkins, Michael; Grayson M (2011). The Eponym Dictionary of Reptiles. Baltimore: Johns Hopkins University Press. xiii + 296 pp. ISBN 978-1-4214-0135-5. (Gloydius blomhoffi, p. 28).
Gloyd HK, Conant R (1990). Snakes of the Agkistrodon Complex: A Monographic Review. Society for the Study of Amphibians and Reptiles. 614 pp. 52 plates. LCCN 89-50342. ISBN 0-916984-20-6. (Agkistrodon blomhoffi complex, pp. 273-309).
Fukuda, Tadashi; Iwaki, Masaaki; Hong, Seung Hwa; Oh, Ho Jung; Wei, Zhu; Morokuma, Kazunori; Ohkuma, Kunio; Dianliang, Lei; Arakawa, Yoshichika; Takahashi, Motohide (2006). "Standardization of Regional Reference for Mamushi (Gloydius blomhoffii ) Antivenom in Japan, Korea, and China". Japanese Journal of Infectious Diseases. 59 (1): 20–4. PMID 16495629.
Hung, Yao-Ching; Sava, Vasyl; Hong, Meng-Yen; Huang, G. Steven (2004). "Inhibitory effects on phospholipase A2 and antivenin activity of melanin extracted from Thea sinensis Linn". Life Sciences. 74 (16): 2037–47. doi:10.1016/j.lfs.2003.09.048. PMID 14967198.
Igari, R; Iseki, K; Abe, S; Syoji, M; Sato, M; Shimomura, K; Hayashida, A; Sugiura, A; Iwashita, Y; Midorikawa, S (2010). 症例報告 マムシ咬傷により複視・眼瞼下垂をきたした1例 [Binocular diplopia and ptosis due to snakebite (Agkistrodon blomhoffi "mamushi")--a case report]. Brain and Nerve (in Japanese). 62 (3): 273–7. PMID 20297733. Archived from the original on 2013-09-05.
Sakurai, Yoshihiko; Shima, Midori; Matsumoto, Tomoko; Takatsuka, Hideo; Nishiya, Katsumi; Kasuda, Shogo; Fujimura, Yoshihiro; Yoshioka, Akira (2003). "Anticoagulant activity of M-LAO, l-amino acid oxidase purified from Agkistrodon halys blomhoffii, through selective inhibition of factor IX". Biochimica et Biophysica Acta (BBA) - Proteins and Proteomics. 1649 (1): 51–57. doi:10.1016/S1570-9639(03)00157-2. PMID 12818190.
Yamazaki, Yasuo; Koike, Hisashi; Sugiyama, Yusuke; Motoyoshi, Kazuko; Wada, Taeko; Hishinuma, Shigeru; Mita, Mitsuo; Morita, Takashi (2002). "Cloning and characterization of novel snake venom proteins that block smooth muscle contraction". European Journal of Biochemistry. 269 (11): 2708–15. doi:10.1046/j.1432-1033.2002.02940.x. PMID 12047379.
Watanabe H, Nagatake T, Matsumoto K, Sakamoto T, Rikitomi N, Hirano E (1992). "Effectiveness of protease inhibition in severe mamushi bite". Procs. XXXIV Annual Meetings of Japan Society of Tropical Medicine, 25–26 Nov 1992, Nagasaki, p.75. Preprint. Also published by same authors as Jpn J Trop Med Hyg, 21(1):39–92, 1993.
Motou K, Yoshida A, Hattori S, Ohno M (2003). "A trial of muscle necrosis prevention by T. flavoviridis venom". Kagoshima University Journal of Medicine 23: 15–24.
Otsuji Y, Irie Y, Ueda H, Yotsueda K, Kitahara T, Yokoyama K, Higashi Y (1978). "A case of acute renal failure caused by Mamushi (Agkistrodon halys) bite". Medical J Kagoshima Univ 30: 129–135. (in Japanese).
Takeshita, T; Yamada, K; Hanada, M; Oda-Ueda, N (2003). "Case report: Extraocular muscle paresis caused by snakebite". Kobe Journal of Medical Sciences. 49 (1–2): 11–5. PMID 12698017.

Nasu, Kaei; Ueda, Tami; Miyakawa, Isao (2004). "Intrauterine Fetal Death Caused by Pit Viper Venom Poisoning in Early Pregnancy". Gynecologic and Obstetric Investigation. 57 (2): 114–6. doi:10.1159/000075676. PMID 14691344. S2CID 39491397.

Further reading

Boie H (1826). "Merkmale eineger japanischer Lurche ". Isis von Oken 19: 203-216. (Trigonocephalus blomhoffii, new species, pp. 214–215). (in German and Latin).

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