Gobiesocidae

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Clingfishes
Temporal range: 23.03-0 Ma Early Miocene–Present
TsuruUU.jpg
Aspasmichthys ciconiae
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Clade: Percomorpha
(unranked): Ovalentaria
Order: Gobiesociformes
Family: Gobiesocidae
Bleeker, 1860
Type species
Gobiesox cephalus
Lacépède, 1800

Clingfishes are ray-finned fishes of the family Gobiesocidae, the only family in the suborder Gobiesocoidei of the order Blenniiformes. [1] These fairly small to very small fishes are widespread in tropical and temperate regions, mostly near the coast, but a few species live in deeper seas or fresh water. Most species shelter in shallow reefs or seagrass beds, clinging to rocks, algae and seagrass leaves with their sucking disc, a structure on their chest. [2] [3]

Contents

They are generally too small to be of interest to fisheries, although the relatively large Sicyases sanguineus regularly is caught as a food fish, [4] and some of the other species occasionally appear in the marine aquarium trade. [2]

Distribution and habitat

Many clingfish live in the intertidal zone and can survive for long periods out of water, as first described for Diplecogaster bimaculata in 1891 Diplecogaster bimaculata (Stefano Guerrieri).jpg
Many clingfish live in the intertidal zone and can survive for long periods out of water, as first described for Diplecogaster bimaculata in 1891
Lepadichthys lineatus and some other clingfish species are associated with crinoids TasujiUUU.jpg
Lepadichthys lineatus and some other clingfish species are associated with crinoids

Clingfishes are primarily found near the shore in the Atlantic, Indian and Pacific Oceans, including marginal seas such as the Mediterranean, Gulf of Mexico, Caribbean and Gulf of California. The greatest species richness is in tropical and warm temperate regions, but the range of a few extends into colder waters, like Diplecogaster bimaculata (north to Norway), Apletodon dentatus , Lepadogaster candolii and L. purpurea (all three north to Scotland; the last formerly mistaken for the mostly Mediterranean L. lepadogaster ), Gobiesox maeandricus (north to Alaska), Gobiesox marmoratus and Sicyases sanguineus (both to southernmost South America), and Gastrocymba quadriradiata (from New Zealand's subantarctic islands). [7] [8] [9] [10] [11]

Clingfishes mainly inhabit shallow rocky reefs and shores, coral reefs, seagrass meadows and algae beds. They often live in places exposed to strong currents and wave action, and some are amphibious. As long as the strongly amphibious, intertidal-living species are kept moist by splashing waves, they can survive for up to three–four days on land, gaining oxygen from the air by the branchial surfaces (gills), skin and perhaps the mouth. [5] [12] [13] At least a few species even tolerate a relatively high degree of water loss when on land. [5]

A relatively small number of species shelter in sea urchins or crinoids. Whether this relationship is obligate (clingfish always with a sea urchin or crinoid) or facultative (clingfish sometimes with a sea urchin or crinoid) varies with species. In some, only young clingfish are obligate and gradually move away as they become adult. [14] [15] [16] Three clingfish species, the Australian Cochleoceps bicolor and C. orientalis, and the warm East Atlantic Diplecogaster tonstricula, are cleaner fish that will cling onto the bodies of larger fish. [2] [17] [18]

Although several species can occur in brackish water, only seven (Gobiesox cephalus, G. fluviatilis, G. fulvus, G. juniperoserrai, G. juradoensis, G. mexicanus and G. potamius) from warmer parts of the Americas are freshwater fish that live in fast-flowing rivers and streams. [19] [20]

Most known clingfish species are from relatively shallow coastal waters, but several inhabit the mesophotic zone and a few even deeper, with Alabes bathys, Gobiesox lanceolatus, Gymnoscyphus ascitus , Kopua kuiteri, K. nuimata and Protogobiesox asymmetricus reported from depths of 300–560 m (980–1,840 ft). [21] [22] Because of their small size and typical habitat, it is however suspected that still-undiscovered deep-water species remain. [21] Even in shallow coastal waters many clingfish are highly cryptic and easily overlooked, mostly staying under cover, although there are species that are active and will swim in the open. [23] As a consequence their abundance is often not well known. Several species are only known from a single or a few specimens. [21] [22] [24] Species that appear uncommon or rare based on standard methods can actually be common if using methods that are more suitable for detecting them. [25] Studies of better-known species have shown that they can be locally abundant. As many as 23 individuals of Lepadogaster lepadogaster have been documented from a single square metre (more than two individuals per square foot). [26] As of 2018, the IUCN has evaluated the conservation status of 84 clingfish species (roughly half the species in the family). The majority of these are considered least concern (not threatened), 17 are considered data deficient (available data prevents an evaluation), 8 considered vulnerable and a single endangered. The vulnerable and endangered species all have small distributions, restricted to islands or a single bay. [27] Three Gobiesox species that are restricted to fresh water in Mexico have not been rated by the IUCN, but are considered threatened by Mexican authorities. [28]

Description

Alabes are eel-like, unlike other clingfish FMIB 45581 Cheilobranchus rufus.jpeg
Alabes are eel-like, unlike other clingfish

Clingfishes are typically small fish, with most species less than 7 cm (2.8 in) in length, [29] and the smallest no more than 1.5 cm (0.6 in). [2] Only a few species can surpass 12 cm (4.7 in) in length and the largest, Chorisochismus dentex and Sicyases sanguineus, both reach up to 30 cm (12 in). [5] [30] Males typically grow larger than females. [3]

Most clingfish species have tapering bodies and flattened heads, appearing somewhat tadpole-like in their overall shape. They lack a swim bladder. The lateral line of clingfish is well developed, but may not extend to the posterior parts of the body. The skin of clingfishes is smooth and scaleless, with a thick layer of protective mucus. [3] In at least Diademichthys lineatus and Lepadichthys frenatus, the mucus production increases if the fish is disturbed. The taste of their mucus is highly bitter to humans and it can kill other fish. This is due to their skin and mucus containing a grammistin-like toxin (the toxin in soapfish, such as Grammistes ). Whether any other clingfish has toxins in its skin or mucus is currently unknown. [6] [31] Another defense appears to be present in a couple of Acyrtus and Arcos species. They have a spine at their gill cover and it appears to be connected to a venom gland. Although the evidence presently is circumstantial, this strongly suggests that the world's smallest venomous fish is Acyrtus artius, which is less than 3 cm (1.2 in) long. [32] [33]

Sucking disc

Sucking disc of Sicyases sanguineus (viewed from below with mouth at the top) Sicyases sanguineus sucker.jpg
Sucking disc of Sicyases sanguineus (viewed from below with mouth at the top)

Clingfish are named for their ability to firmly attach themselves to various surfaces, even in strong water currents or when battered by waves. This ability is aided by their sucking disc, which is located on the underside at the chest and is formed primarily by modified pelvic fins and adjacent tissue. [3] [5] [13] [29] In some species it is divided in two, resulting in a larger front and a smaller rear sucking disc. [3] The sucking disc is covered in tiny hexagons and each of these consists of many microscopic hair-like structures (setae). This is similar to the structures that allow geckos to cling to walls. The sucking disc can be remarkably strong, in some species able to lift as much as 300 times the weight of the clingfish. [13] Gobies (family Gobiidae) can have a similar sucking disc, but unlike that family the single dorsal fin in clingfish is not spiny. [3] In a few clingfish species the disc is reduced or even absent, notably Alabes, which are quite eel-like in their shape and aptly named shore-eels. [2] [17] The sucking disc is also reduced in some deep-water clingfish species. [13]

Colours

Gobiesox rhessodon.jpg
Lepadogaster purpurea (Cornwall, England).jpg
HashinagaUU.jpg
Eastern Cleaner Clingfish-Cochleoceps orientalis (22695734536).jpg

Most clingfish species have a cryptic colouration, often brown, grey, whitish, black, reddish or green shades, and in some cases they can rapidly change colour to match their background. [3] [34] [35]

Species of deep water are often orange-red (these long wave-length colours are the first that disappear with depth, making them suitable for camouflage). [22] Diademichthys lineatus, Discotrema species, Lepadichthys caritus and L. lineatus are strongly banded, which may function as a disruptive pattern when among sea urchin spines or crinoid arms, but may also be warning colours, as some members of these genera have poisonous skin and mucus (it is unknown if all of them are poisonous). [6] [14] [31] There are species with colours or patterns that are unsuitable for camouflage. Although Lepadogaster purpurea overall is cryptic, it has a pair of distinct large eyespots on the top of its head. [10] Cochleoceps bicolor, C. orientalis and Diplecogaster tonstricula are yellow to red with fine bluish lines. These three are cleaner fish. [17] [18]

Feeding

Feeding varies depending on exact clingfish species. Most primarily feed on tiny crustaceans (such as amphipods, copepods, isopods, mysids, ostracods and shrimp) or gastropods (limpets and other sea snails). Other small animals that have been recorded in their diet include chitons, bivalves, medium-small crustacean like crabs and barnacles, sea urchins, worms, insect larvae, fish and fish eggs. [3] [4] [35] [36] In some species, cannibalism where a large clingfish eats a smaller clingfish is not uncommon. [35] [37]

Limpets and other shelled invertebrates are well-protected and often strongly attached to the rock surface. Clingfish species that feed extensively on them have developed specialized teeth and techniques to dislodge them. This includes rapidly inserting their relatively large, fang-like front teeth under the edge of the prey to flip it, or jamming the teeth on or under the shell's edge to make a small break. [12] [13] [30] However, the teeth of clingfish vary extensively depending on species. [38] [39] In the opposite extreme of the species with relatively few large teeth is Nettorhamphos radula . This species has 1,800–2,300 microscopic teeth (about ten times more than known from any other clingfish), but its feeding behavior is unknown. [24] [38]

Three clingfish species, Cochleoceps bicolor, C. orientalis and Diplecogaster tonstricula, have become cleaner fish. Large fish approach them and allow the small clingfish onto their body where the clingfish eats tiny parasites. [17] [18] In contrast to this mutualistic relationship, certain clingfish species that live among the spines of sea urchins appear to be part of a more varied relationship. It can be either commensal (the clingfish gains protection from the sea urchin spines, but apparently neither benefits nor is a disadvantage to the sea urchin) or parasitic (the clingfish gains protection, and eats tube feet and pedicellaria from its sea urchin host). [15] [16] [40]

No clingfish species is known to be exclusively herbivorous, but some are omnivorous and will feed extensively on a range of algae (brown, green and red), [4] while other, more strictly carnivorous species may ingest plant material incidentally. [35]

Classification and taxonomy

The classification of the clingfishes varies. FishBase places Gobiesocidae as the only family in the order Gobiesociformes, under the superorder Paracanthopterygii; [41] whereas ITIS place them in the suborder Gobiesocoidei of the order Perciformes, under superorder Acanthopterygii. ITIS lists Gobiesociformes as invalid. [42] The 5th edition of Fishes of the World places the Gobiesociiformes in the clade Percomorpha as part of the series Ovalentaria. [43]

Mostly being very small and often cryptic, new species are regularly discovered and described. A major authoritative work on the family is a monograph that was published in 1955 by J.C. Briggs, [44] but in the half century after its publication, up until 2006, fifty-six new clingfish species were described, or on average more than one per year. [6] This pattern with regular descriptions of new species—and even new genera—has continued since then. [38] [45] [46] [47] As of 2020, there are 182 recognized clingfish species. [48]

Subfamilies and genera

Chorisochismus dentex is the only member of its genus and the largest clingfish species Rocksucker at Blousteen ridge DSC09449.JPG
Chorisochismus dentex is the only member of its genus and the largest clingfish species
The Connemara clingfish is typically referred to as Lepadogaster candolii, but genetic studies indicate that this makes the genus Lepadogaster polyphyletic Lepadogaster candolii Rab.JPG
The Connemara clingfish is typically referred to as Lepadogaster candolii, but genetic studies indicate that this makes the genus Lepadogaster polyphyletic
Lepadogaster lepadogaster (shown) and L. purpurea were formerly considered a single species. Both inhabit the Mediterranean and the East Atlantic, but only L. purpurea ranges north to the British Isles Lepadogaster lepadogaster 1.jpg
Lepadogaster lepadogaster (shown) and L. purpurea were formerly considered a single species. Both inhabit the Mediterranean and the East Atlantic, but only L. purpurea ranges north to the British Isles

Subfamilies and genera. The delimination of the subfamilies, and to some extent the genera, is not fully resolved. [23] [47] The 5th edition of Fishes of the World recognises only two subfamilies, Cheilobranchinae and Gobiesocinae. [43] Fishbase does list a third subfamily, the monotypic Protogobiesocinae which contains a single species Protogobiesox asymmetricus , this species having been described in 2016. [49] In 2020 the systematics of Gobiesocidae was reviewed and nine subfamilies were proposed: Cheilobranchinae, Chorisochisminae, Diademichthyinae, Diplocrepinae, Haplocylicinae, Gobiesocinae, Lepadogastrinae, Protogobiesocinae, and Trachelochisminae. [48]

Subfamily Cheilobranchinae

Subfamily Chorisochisminae

Subfamily Diademichthyinae

Subfamily Diplocrepinae

Subfamily Gobiesocinae

Subfamily Haplocylicinae

Subfamily Lepadogastrinae

Subfamily Protogobiesocinae

Subfamily Trachelochisminae

Incertae Sedis

Related Research Articles

<span class="mw-page-title-main">New Zealand urchin clingfish</span> Species of fish

The New Zealand urchin clingfish is a clingfish. It is found around New Zealand wherever sea urchins are present. Its length is between 2 and 3 cm.

<i>Dellichthys</i> Genus of fishes

Dellichthys is a small genus of clingfishes from the family Gobiesocidae which are endemic to New Zealand. It had been regarded as a monotypic genus but a second species was described in 2018.

<span class="mw-page-title-main">Shore clingfish</span> Species of fish

The shore clingfish is a clingfish of the family Gobiesocidae. It is found in the Mediterranean Sea and adjacent Atlantic Ocean north to Galicia, Spain.

<i>Diademichthys lineatus</i> Species of fish

Diademichthys lineatus, commonly known as the long-snout clingfish or urchin clingfish, is a species of marine fish in the family Gobiesocidae.

<span class="mw-page-title-main">Northern clingfish</span> Species of fish

The northern clingfish is a species of saltwater fish. It is a member of the family Gobiesocidae of order Gobiesociformes. It is native to the Pacific coast of North America from Baja California north to Revillagigedo Island Alaska. It is commonly found in the intertidal zone clinging to the underside of rocks by small hairs akin to those on a gecko's feet on the basal portions of the pelvic and pectoral fins. This species was described by Charles Frédéric Girard in 1858 from specimens collected at San Luis Obispo in California, Girard had originally named it as Lepadogaster reticulatus in 1854 but this name was preoccupied by Lepadogaster reticulatus Risso, 1810.

<i>Sicyases sanguineus</i> Species of fish

Sicyases sanguineus is a species of amphibious marine clingfish in the family Gobiesocidae. It lives in the Southeast Pacific along the entire coast of Chile and southern Peru.

<i>Acyrtus</i> Genus of fishes

Acyrtus is a genus of clingfishes found in the western Atlantic Ocean.

Derilissus is a genus of clingfishes belonging to the family Gobiesocinae found in the western Atlantic Ocean. This family of fish is identified by their appearance as small fish with sucking discs which allow them to attach themselves to various surfaces. Derilissus differs from other genera due to its attached gill membranes.

<i>Gobiesox</i> Genus of fishes

Gobiesox is a genus of clingfishes found in the Americas, including offshore islands. Most species inhabit coastal marine and brackish waters, but G. lanceolatus is a deep-water species found at a depth of around 300 m (980 ft), and seven species are from fast-flowing rivers and streams. These seven are the only known freshwater clingfish.

<span class="mw-page-title-main">Blunt-snouted clingfish</span> Species of fish

The blunt-snouted clingfish is a species of clingfish found along the western Mediterranean Sea coasts from Spain to Italy. This species grows to a length of 5 centimetres (2.0 in) TL. The blunt-nosed clingfish is a little known species of shallow water along the littoral of the northern Mediterranean from Alicante to Sicily. Its range was formerly thought to extend to Israel and Syria in the east; however this was actually several closely related cryptic species. It can survive out of the water and occurs only among intertidal pebbles and sand.

<i>Gouania</i> (fish) Genus of fish

Gouania is a genus of clingfishes endemic to the Mediterranean Sea that contains at least 5 cryptobenthic species. The members of this genus are the only vertebrates to inhabit European intertidal gravel beaches and they have developed many adaptations to survive in this extremely harsh environment. They are a type of Clingfish meaning they form part of the family Gobiesocidae. All species of clingfish have a thoracic adhesive disc that allows them to hold on to both smooth and rough surfaces alike. This adaptation enables Gouania to survive the harsh conditions of gravel beaches, they are amongst the only fish taxa adapted to this habitat.

Gymnoscyphus ascitus is a small species of clingfish found in the Caribbean Sea at depths of 231 to 318 m (758–1,043 ft). This species is the only known member of the genus Gymnoscyphus.

<i>Kopua</i> Genus of fishes

Kopua is a genus of clingfishes found in the Pacific Ocean.

<i>Lepadichthys</i> Genus of fishes

Lepadichthys is a genus of clingfishes native to the Indian and Pacific Oceans.

<i>Lepadogaster</i> Genus of fishes

Lepadogaster is a genus of clingfishes native to the eastern Atlantic Ocean extending into the Mediterranean Sea. Lepadogaster belongs to class Actinopterygii. This means that they share many of the same characteristics as eels, ray-finned fish, and sea horses to name a few. The main characteristic of all of them though is having fin rays. These fin rays are made of webbed skin and are attached to portions of the body that connect fins to the bones. Lepadogaster species have a distinct difference in the formation of their dorsal and anal fins. While most other ray-finned fish spines, branched fin-rays, and middle radials, Lepadogaster species do not have these. Instead, they have cartilage in place of the mentioned features. These clingfish are mainly found near the rocky coasts and inside intertidal zones. Lepadogaster is known mostly as a clingfish, meaning that it spends most of its time attached to the surface of rocks.

Pherallodiscus is a genus of clingfishes native to the central eastern Pacific Ocean along the coast of Mexico. Based on genetic studies the genus should be merged into Gobiesox.

<i>Lepadogaster candolii</i> Species of fish

Lepadogaster candolii, common name Connemarra clingfish, is a species of fish in the genus Lepadogaster. It occurs in the Eastern Atlantic from the British Isles south to Madeira and the Canary Islands and into the western Mediterranean and the Black Sea. The specific name candolii honours the Swiss botanist Augustin Pyramus de Candolle (1778–1841) and has various spellings: candolii, candolei, candollei, and decandollii, but only the first one is correct. Some workers have found that L. candolii is not closely related to the other two species in the genus Lepadogaster and have proposed the placing of this species in the revived monotypic genus Mirbelia Canestrini, 1864, at least until more definitive taxonomic studies can be undertaken.

Diplecogaster tonstricula, commonly known as the Eastern Atlantic cleaner clingfish, is a species of clingfish from the family Gobiesocidae, which is found in the tropical eastern North Atlantic Ocean. It has been observed cleaning larger species of fish.

<i>Flexor incus</i> Species of fish

Flexor is a genus of clingfish from the family Gobiesocidae. It is a monotypic genus, the sole member of which Flexor incus was described in 2018 by Kevin W. Conway, Andrew L. Stewart and Adam P. Summers from type specimens collected on Raoul Island and L'Esperance Rock in the Kermadec Islands. An unknown species of clingfish was known to occur in the Kermadecs and it had been provisionally assigned to the genus Aspasmogaster. When examined, the specimens proved to belong to a new genus which they name Flexor, in reference to the flexibility of clingfishes, and incus, a reference to the anvil-like shape of the type locality, Raoul Island.

<i>Opeatogenys gracilis</i> Species of fish

Opeatogenys gracilis is a species of clingfish from the family Gobiesocidae which is found in the Mediterranean Sea and in the eastern Atlantic Ocean. Suggested common names for this species are the pygmy clingfish and the seagrass clingfish.

References

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  9. ↑ Froese, Rainer; Pauly, Daniel (eds.). "Gastrocymba quadriradiata". FishBase . October 2018 version.
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  25. ↑ Craig, M.T.; Williams, J.T. (2015). "Arcos nudus". IUCN Red List of Threatened Species . 2015: e.T185939A1792278. doi: 10.2305/IUCN.UK.2015-2.RLTS.T185939A1792278.en .
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  27. ↑ The IUCN Red List of Threatened Species (12 October 2018). "Gobiesocidae" . Retrieved 12 October 2018.
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  29. 1 2 Froese, Rainer; Pauly, Daniel (eds.). "Family Gobiesocidae". FishBase . October 2012 version.
  30. 1 2 3 Stobbs, R.E. (1980). "Feeding Habits of the Giant Clingfish Chorisochismus dentex (Pisces: Gobiesocidae)". South African Journal of Zoology. 15 (3): 146–149. doi: 10.1080/02541858.1980.11447702 .
  31. 1 2 Hori, K.; N. Fusetani; K. Hashimoto; K. Aida; J.E. Randall (1979). "Occurrence of a grammistin-like mucous toxin in the clingfish Diademichthys lineatus". Toxicon. 17 (4): 418–424. Bibcode:1979Txcn...17..418H. doi:10.1016/0041-0101(79)90271-X. PMID   494325.
  32. ↑ Conway, K.W.; C. Baldwin; M.D. White (2014). "Cryptic Diversity and Venom Glands in Western Atlantic Clingfishes of the Genus Acyrtus (Teleostei: Gobiesocidae)". PLOS ONE. 9 (5): e97664. Bibcode:2014PLoSO...997664C. doi: 10.1371/journal.pone.0097664 . PMC   4019652 . PMID   24825326.
  33. ↑ Keartes, S. (22 May 2014). "Could This be the World's Smallest Venomous Fish?". Earth Touch News. Retrieved 12 October 2018.
  34. ↑ Briggs, J.C.; Hutchins, J.B. (1998). Paxton, J.R.; Eschmeyer, W.N. (eds.). Encyclopedia of Fishes. San Diego: Academic Press. pp. 142–143. ISBN   978-0-12-547665-2.
  35. 1 2 3 4 Pires, T.H.S.; F.Z. Gibran (2011). "Intertidal life: field observations on the clingfish Gobiesox barbatulus in southeastern Brazil" (PDF). Neotrop. Ichthyol. 9 (1): 233–240. doi: 10.1590/S1679-62252011005000001 .
  36. ↑ Hirayama, S.; T. Shiiba; Y. Sakai; H. Hashimoto; K. Gushima (2005). "Fish-egg predation by the small clingfish Pherallodichthys meshimaensis (Gobiesocidae) on the shallow reefs of Kuchierabu-jima Island, southern Japan". Environmental Biology of Fishes. 73 (3): 237–242. Bibcode:2005EnvBF..73..237H. doi:10.1007/s10641-005-2260-2. S2CID   31859123.
  37. ↑ Johnson, C.R. (1970). "Notes on the Intertidal Life History of the Northern Clingfish, Gobiesox maeandricus (Girard)". The American Midland Naturalist. 83 (2): 625–627. doi:10.2307/2423966. JSTOR   2423966.
  38. 1 2 3 Conway, K.W.; G.I. Moore; A.P. Summers (2017). "A New Genus and Species of Clingfish (Teleostei: Gobiesocidae) from Western Australia". Copeia. 105 (1): 128–140. doi:10.1643/CI-16-560. S2CID   90595940.
  39. ↑ Conway, K.W.; N.G. Bertrand; Z. Browning; Z.W. Lancon; F.J. Clubb, Jr. (2015). "Heterodonty in the New World: An SEM Investigation of Oral Jaw Dentition in the Clingfishes of the Subfamily Gobiesocinae (Teleostei: Gobiesocidae)". Copeia. 103 (4): 973–998. doi:10.1643/OT-15-234. S2CID   86305371.
  40. ↑ Russell, B.C. (1983). "The food and feeding habits of rocky reef fish of north-eastern New Zealand". New Zealand Journal of Marine and Freshwater Research. 17 (2): 121–145. Bibcode:1983NZJMF..17..121R. doi:10.1080/00288330.1983.9515991.
  41. ↑ Froese, Rainer; Pauly, Daniel (eds.). "Order Gobiesociformes". FishBase . october 2012 version.
  42. ↑ "Gobiesociformes". Integrated Taxonomic Information System . Retrieved 5 February 2008.
  43. 1 2 J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. p. 351. ISBN   978-1-118-34233-6. Archived from the original on 8 April 2019. Retrieved 2 June 2019.
  44. ↑ Briggs, J.C. (1955). "A monograph of the clingfishes (Order Xenopterygii)". Stanford Ichthyological Bulletin. 6: 1–224.
  45. ↑ Fricke, R. (2014). "Unguitrema nigrum, a new genus and species of clingfish (Teleostei: Gobiesocidae) from Madang, Papua New Guinea" (PDF). Journal of the Ocean Science Foundation. 13: 35–42.
  46. ↑ Fricke R., Chen J.-N., Chen W.-J. (2016). "New case of lateral asymmetry in fishes: A new subfamily, genus and species of deep water clingfishes from Papua New Guinea, western Pacific Ocean" (PDF). Comptes Rendus Biologies. 340 (1): 47–62. doi: 10.1016/j.crvi.2016.11.002 . PMID   27979388.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  47. 1 2 Conway, K.W.; A.L. Stewart; A.P. Summers (2018). "A new genus and species of clingfish from the Rangitāhua Kermadec Islands of New Zealand (Teleostei, Gobiesocidae)". ZooKeys (786): 75–104. Bibcode:2018ZooK..786...75C. doi: 10.3897/zookeys.786.28539 . PMC   6168618 . PMID   30283237.
  48. 1 2 Kevin W. Conway; Cragen D. King; Adam P. Summers; Daemin Kim; Philip A. Hastings; Glenn I. Moore; Samuel P. Iglésias; Mark V. Erdmann; Carole C. Baldwin; Graham Short; Kyoji Fujiwara; Thomas Trnski; Gary Voelker; Lukas Rüber (23 December 2020). "Molecular Phylogenetics of the Clingfishes (Teleostei: Gobiesocidae)—Implications for Classification" (PDF). Ichthyology & Herpetology. 108 (4): 886–906. doi: 10.1643/CI2020054 . eISSN   2766-1520. ISSN   2766-1512. S2CID   230507140.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  49. ↑ Froese, Rainer; Pauly, Daniel (eds.). "Species in genus Protogobiesox". FishBase . April 2019 version.
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