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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Superclassis/Classis: Actinopterygii
Classis/Subclassis: Actinopteri
Subclassis/Infraclassis: Neopterygii
Infraclassis: Teleostei
Megacohors: Osteoglossocephalai
Supercohors: Clupeocephala
Cohors: Euteleosteomorpha
Subcohors: Neoteleostei
Infracohors: Eurypterygia
Sectio: Ctenosquamata
Subsectio: Acanthomorphata
Divisio/Superordo: Acanthopterygii
Ordo: Scorpaeniformes
Subordo: Scorpaenoidei

Familia: Synanceiidae
Genus: Inimicus
Species (10): I. brachyrhynchus – I. caledonicus – I. cuvieri – I. didactylus – I. filamentosus – I. gruzovi – I. japonicus – I. joubini – I. sinensis – I. smirnovi
Name

Inimicus Jordan & Starks, 1904: 158

Type species: Pelor japonicum Cuvier, 1829, by original designation.
References
Primary references

Jordan, D.S. & Starks, E.C. 1904. A review of the scorpaenoid fishes of Japan. Proceedings of the United States National Museum 27(1351): 91–175, pls. 1–2.
Inaba, T. & Motomua, H. 2018. Review of the Indo-West Pacific genus Inimicus (Synanceiidae: Choridactylinae). Zootaxa 4482(1): 52–90. DOI: 10.11646/zootaxa.4482.1.2 Paywall Reference page.

Inimicus is a genus of marine ray-finned fishes, it is one of two genera in the tribe Choridactylini, one of the three tribes which are classified within the subfamily Synanceiinae within the family Scorpaenidae, the scorpionfishes and their relatives. These venomous, benthic fishes are found on sandy or silty substrates of lagoon and seaward reefs, in coastal regions of tropical oceans. The ten described species are collectively known by various common names, including ghoul, goblinfish, sea goblin, spiny devilfish, stinger, and stingfish.

Taxonomy

Inimicus was first described as a genus in 1904 by the American ichthyologists David Starr Jordan and Edwin Chapin Starks with the type species designated as Pelor japonicum,[1] which had been described by Georges Cuvier in 1829 from the seas of China and Japan.[2] It is one of two genera in the tribe Choridactylini within the subfamily Synanceiinae of the family Scorpaenidae.[3] However, some authorities classify this taxon as a subfamily Choridactylinae within the family Synanceiidae.[1] The genus name inimicus is Latin for "foe" or "enemy", Fishermen feared these fishes because of their venomous spines.[4]
Species

The members of the genus Inimicus are nearly identical in appearance and behavior, and often confused with one another. There are currently 10 recognized species in this genus:[5]
Species[6] Common name[6] Habitat[6] Distribution[6] IUCN status[7]
Inimicus brachyrhynchus
(Bleeker, 1874)[8] Marine; tropical; reef-dwelling; demersal Western Central Pacific NE*
Inimicus caledonicus
(Sauvage, 1878)[9] Chinese ghoul, Caledonian stinger, demon stinger Marine; tropical; reef-dwelling; demersal Eastern Indian Ocean: Andaman and Nicobar islands. Western Pacific: Australia, Papua New Guinea and New Caledonia. LC
Inimicus cuvieri
(J. E. Gray, 1835)[10] Marine; tropical; reef-dwelling; demersal Western Pacific: South China Sea NE
Inimicus didactylus
(Pallas, 1769)[11] Goblinfish, popeyed sea goblin, demon stinger, devil stinger, longsnout stinger, spiny devilfish, bearded ghoul Marine; tropical; reef-dwelling; demersal Eastern Indian and Western Pacific oceans NE
Inimicus filamentosus
(G. Cuvier, 1829)[12] Filament-finned stinger, barred ghoul, two-stick stingfish, devil scorpionfish Marine; tropical; reef-dwelling; demersal Western Indian Ocean: Red Sea and East Africa to Maldives LC
Inimicus gruzovi
Mandritsa, 1991[13] Marine; tropical; reef-dwelling; demersal Western Central Pacific NE
Inimicus japonicus
(G. Cuvier, 1829)[14] Devil stinger Marine; tropical; reef-dwelling; demersal Indo-West Pacific: Japan and East China Sea NE
Inimicus joubini
(Chevey, 1927)[15] Marine; tropical; reef-dwelling; demersal Northwest Pacific: Japan and Vietnam NE
Inimicus sinensis
(Valenciennes, 1833)[16] Spotted ghoul, spotted stonefish Marine; tropical; reef-dwelling; demersal LC
Inimicus smirnovi
Mandritsa, 1990[17] Marine; tropical; reef-dwelling; demersal Western Central Pacific NE

NE: Not Evaluated

Species no longer recognized:

Inimicus barbatus (De Vis, 1884) is a jr. synonym of Inimicus caledonicus[18]
Inimicus dactylus (Cornic, 1987) is a jr. synonym of Inimicus filamentosus[19]

Geographic distribution

Members of the genus Inimicus are distributed mainly in warm tropical waters in the coastal regions of Indo-Pacific oceans. Their range does however extend a little into the subtropical zone. The waters of the Red Sea off the coast of Egypt appear to mark the westernmost limit of their range, while specimens have been reported as far to the east as New Caledonia. The northern coast of New South Wales, Australia marks the southernmost extent of their range, which extends as far to the north as Japan's Aomori, the northernmost prefecture of Japan's main island.[20][21] Inimicus are benthic fishes, living mainly on the bottom of mangrove swamps and coral reefs, at depths between 5 and 450 meters.[22][23]
Description

Adults are typically 13-25 centimeters in length, and can weigh up to 480 grams. The body color can be a dull yellow, gray, brown, or rust in color with light blotches, and very similar to that of the surrounding sandy or coral seabed in which they are found. This coloration acts as a camouflage which renders them extremely difficult to detect in their natural habitat. The skin is without scales except along the lateral line, and is covered with venomous spines and wartlike glands which give it a knobby appearance. The head is flattened, depressed and concave. The eyes, mouth and nostrils project upwards and outwards from the dorsal aspect of the head. Sexual dimorphism is not believed to occur in this genus.[citation needed]

Fin morphology:

dorsal fin: composed of 15 to 17 spines and 7 to 9 soft rays.[22][23][24]
caudal fin: composed of 2-4 spines and 4-14 soft rays, with dark bands at basal and subterminal positions.
pelvic fin: composed of one spine and 3-5 soft rays.
pectoral fin: composed of 10-12 rays. The two most caudal rays of each pectoral fin are detached from the rest of the fin, and angled in a ventral direction. The fish employ these two rays to prop up the forward part of their body, as well as to "walk" along the bottom of the substrate.[25][26][27][28] The ventral surface of the pectoral fins bears broad black bands containing smaller, lighter spots at the basal and distal ends. In I. filamentosus, these bands are attenuated, while the bands of I. sinensis have yellow spots on them. This is a key feature for distinguishing the two species, which are otherwise nearly identical.[26]

Behavior

Inimicus are piscivorous ambush predators. They are nocturnal and typically lie partially buried on the sea floor or on a coral head during the day, covering themselves with sand and other debris to further camouflage themselves. They have no known natural predators. When threatened, they spread their brilliantly colored pectoral and caudal fins as a warning. Once dug in, they are very reluctant to leave their hiding places. When they do move, they display an unusual mechanism of subcarangiform locomotion---they crawl slowly along the seabed, employing the four lower rays (two on each side) of their pectoral fins as legs.[22][23][25][26][27][28]

The paired pectoral fins of these fishes are a remarkable example of their adaptation to life in a benthic environment. No longer useful or necessary for aiding the animal in maneuvering within the water column, the fins have taken on a number of other functions useful to life as a demersal fish. Among these include probing for food items, propping the forward part of the body away from the bottom, and the aforementioned subcarangiform locomotion.[25]

Inimicus is not the only fish that demonstrate this type of ambulation; it has been extensively described in other related benthic Scorpaeniformes fish such as the Sea robin, Flying gurnards, and the Tub Gurnard, Chelidonichthys lucerna. This type of locomotion, based on voluntary and coordinated movements of paired pectoral fins, is believed by some to be a precursor to the later development of similar ambulation in terrestrial vertebrates.[29]
Relevance to humans

Like all known members of the family Synanceiidae, all members of the genus Inimicus possess a complex and extremely potent venom. It is stored in glands at the bases of needle-like spines in their dorsal fins. Upon contact with the dorsal fin, the fish can deliver a very painful, potentially fatal, sting. The venom consists of a mixture of proteolytic enzymes, including stonustoxin (a hemotoxin), trachynilysin (a neurotoxin), and cardioleputin (a cardiotoxin).[citation needed] Envenomation results severe and immediate local pain, sometimes followed by shock, paralysis, tissue necrosis, and even death.[citation needed] The genus name means enemy in Latin.[30]

Despite the obvious risks, one species of Inimicus, I. japonicum, is commercially cultured in Japan. It is used as a food fish there, and it also has applications in Chinese medicine.[citation needed]
Treatment of envenomation
Envenomation by Inimicus species is characterized by immediate and severe local pain. Medical aid must be sought at the earliest opportunity after envenomation. Recommended first aid treatment includes immersion of the affected area in hot water.[31] Immersing the injured area in water at a temperature of at least 45 °C (113 °F) can partially denature the proteolytic enzymes in the venom. Some relief can also be obtained by infiltrating the envenomation site with a local anesthetic. For more extreme cases, an intramuscular injection of a specific horse-derived antivenom can be lifesaving.[32] Tetanus toxoid vaccine should also be administered, if indicated. Surviving victims often suffer localized tissue necrosis and nerve damage, leading to atrophy of adjoining muscle tissues.

References

Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Genera in the family Choridactylinae". Catalog of Fishes. California Academy of Sciences. Retrieved 6 April 2022.
Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Species in the genus Inimicus". Catalog of Fishes. California Academy of Sciences. Retrieved 6 April 2022.
J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 468–475. ISBN 978-1-118-34233-6.
Christopher Scharpf & Kenneth J. Lazara, eds. (10 March 2022). "Order Perciformes (Part 10): Suborder Scorpaenoidei: Families Apistidae, Tetrarogidae, Synanceiidae, Aploacrinidae, Perryenidae, Eschmeyeridae, Pataceidae, Gnathanacanthidae, Congiopodidae and Zanclorhynchidae". The ETYFish Project Fish Name Etymology Database. Christopher Scharpf and Kenneth J. Lazara. Retrieved 6 April 2022.
Froese, Rainer and Pauly, Daniel, eds. (2012). Species of Inimicus in FishBase. December 2012 version.
Froese, R. and D. Pauly. Editors. 2010.FishBase. World Wide Web electronic publication. Fish Identification at FishBase, version (01/2010).Genus Inimicus. Accessed 23 March 2010.
IUCN 2010. The IUCN Red List of Threatened Species Archived 2014-06-27 at the Wayback Machine. Version 2010.1. Accessed 21 March 2010.
"Inimicus brachyrhynchus (Bleeker, 1874)". Encyclopedia of Life, available from "http://www.eol.org/pages/217784". Accessed 21 Mar 2010.
"Inimicus caledonicus (Sauvage, 1878)". Encyclopedia of Life, available from "http://www.eol.org/pages/218118". Accessed 21 Mar 2010.
"Inimicus cuvieri (Gray, 1835)". Encyclopedia of Life, available from "http://www.eol.org/pages/206555". Accessed 21 Mar 2010.
"Inimicus didactylus (Pallas, 1769)". Encyclopedia of Life, available from "http://www.eol.org/pages/211677". Accessed 21 Mar 2010.
"Inimicus filamentosus (Cuvier, 1829)". Encyclopedia of Life, available from "http://www.eol.org/pages/204576". Accessed 21 Mar 2010.
"Inimicus gruzovi Mandrytsa, 1991". Encyclopedia of Life, available from "http://www.eol.org/pages/217782". Accessed 21 Mar 2010.
"Inimicus japonicus (Cuvier, 1829)". Encyclopedia of Life, available from "http://www.eol.org/pages/225253". Accessed 21 Mar 2010.
"Inimicus joubini (Chevey, 1927)". Encyclopedia of Life, available from "http://www.eol.org/pages/583268". Accessed 21 Mar 2010.
"Inimicus sinensis (Valenciennes, 1833)". Encyclopedia of Life, available from "http://www.eol.org/pages/206520". Accessed 21 Mar 2010.
"Inimicus smirnovi Mandrytsa, 1990". Encyclopedia of Life, available from "http://www.eol.org/pages/217783". Accessed 21 Mar 2010.
Eschmeyer, W.N. (ed.) 2004 Catalog of fishes. Updated database version of January 2004. Catalog databases as made available to FishBase in January 2004.
Fricke, R. 1999 Fishes of the Mascarene Islands (Réunion, Mauritius, Rodriguez): an annotated checklist, with descriptions of new species. Koeltz Scientific Books, Koenigstein, Theses Zoologicae, Vol. 31: 759 p.
Wheeler, Alwyne C. (1985). The world encyclopedia of fishes. London: Macdonald & Company.
"Bulletin of Aomori Prefectural Fisheries Research Center" (PDF). Aomori Prefectural Fisheries Research Center. 2004. Retrieved 2016-04-07.
Munro, Ian Stafford Ross (1967). The fishes of New Guinea. Port Moresby, Papua and New Guinea: Department of Agriculture, Stock and Fisheries.
Myers, Robert F. (1999). Micronesian reef fishes : a field guide for divers and aquarists. Barrigada, Territory of Guam, USA: Coral Graphics.
Mandritsa, S.A. (1991). "New species of the genus Inimicus (Scorpaeniformes, Synanceiidae) from the Coral sea". J. Ichthyol. 31 (2): 76–79.
William A. Gosline (July 1994). "Function and structure in the paired fins of scorpaeniform fishes". Journal Environmental Biology of Fishes. 40 (3): 219–226. doi:10.1007/BF00002508. hdl:2027.42/42637. S2CID 30229791.
World Database of Marine Species: Spiny devil fish Archived 2012-03-04 at the Wayback Machine. Accessed 03-22-2010.
Scott Michael (Winter 2001). "Speak of the devil: fish in the genus Inimicus" (PDF). SeaScope. 18. Archived from the original (PDF) on 2011-07-13. Retrieved 2010-03-27.
WetWebMedia.com: The Ghoulfish/Scorpion/Stonefishes of the Subfamily Choridactylinae (Inimicinae), by Bob Fenner. Accessed 03-27-2010.
Marc Jamon; Sabine Renous; Jean Pierre Gasc; Vincent Bels; John Davenport (July 2007). "Evidence of force exchanges during the six-legged walking of the bottom-dwelling fish, Chelidonichthys lucerna". Journal of Experimental Zoology Part A: Ecological Genetics and Physiology. 307A (9): 542–547. doi:10.1002/jez.401. PMID 17620306. Retrieved 2010-03-22.[dead link]
"Inimicus sinensis, Spotted ghoul". FishBase. Retrieved 24 March 2021.
Marine Bites and Stings Archived 2014-03-21 at the Wayback Machine Dr Mark Little

Taylor, G. (2000). "Toxic fish spine injury: Lessons from 11 years experience". South Pacific Underwater Medicine Society Journal. 30 (1). ISSN 0813-1988. OCLC 16986801. Retrieved 2010-03-22.

Further reading
William N. Eschmeyer; Kaza V. Rama-Rao; Leon E. Hallacher (1979). Fishes of the Scorpionfish subfamily Choridactylinae from the Western Pacific and the Indian Ocean. San Francisco: California Academy of Sciences, Proceedings of the California Academy of Sciences, vol. 41(21). pp. 475–500.
Rudie H. Kuiter (1996). Guide to Sea Fishes of Australia. Frenchs Forest, NSW: New Holland.
John E. Randall; Gerald R. Allen; Roger C. Steene (1997). Fishes of the Great Barrier Reef and Coral Sea. Bathurst, NSW: Crawford House Press.
Ewald Lieske; Robert Myers (1994). Coral Reef Fishes: Caribbean, Indian Ocean, and Pacific Ocean Including the Red Sea (Collins Pocket Guide). Harpercollins Publishers

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