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Mudskipper
Periophthalmus barbarus mudskippers in The Gambia
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Order:	Gobiiformes
Family:	Oxudercidae
Subfamily:	Oxudercinae Günther 1861[1]
Genera
Apocryptes Apocryptodon Boleophthalmus Oxuderces Parapocryptes Periophthalmodon Periophthalmus Pseudapocryptes Scartelaos Zappa
Synonyms
Periophthalminae Periophthalmidae

Mudskippers are amphibious fish, presently included in the family Oxudercidae,^[2] in the subfamily Oxudercinae. There are 32 living species of mudskipper. They use their pectoral fins and pelvic fins to walk on land ^[3][4] They typically live in intertidal habitats, and exhibit unique adaptations to this environment that are not found in most intertidal fishes, which typically survive the retreat of the tide by hiding under wet seaweed or in tide pools.^[5] Mudskippers are quite active when out of water, feeding and interacting with one another, for example, to defend their territories and court potential partners. They are found in tropical, subtropical, and temperate regions, including the Indo-Pacific and the Atlantic coast of Africa.

Oxudercinae is sometimes classified within the family Gobiidae (gobies).^[6] Recent molecular studies do not support this classification, as oxudercine gobies appear to be paraphyletic relative to amblyopine gobies (Gobiidae: Amblyopinae), thus being included in a distinct "Periophthalmus lineage", together with amblyopines.^[7] Mudskippers can be defined as oxudercine gobies that are "fully terrestrial for some portion of the daily cycle" (character 24 in Murdy, 1989^[6]). This would define the species of the genera Boleophthalmus, Periophthalmodon, Periophthalmus, and Scartelaos as "mudskippers". However, field observations of Zappa confluentus suggest that this monotypic genus should be included in the definition.^[8]

Adaptations

Periophthalmus gracilis (from Malaysia to northern Australia)

Periophthalmus barbarus (from western Africa)

Compared with fully aquatic gobies, these specialized fish present a range of peculiar anatomical and ethological adaptations that allow them to move effectively on land as well as in the water.^[4][9] As their name implies, these fish use their fins to move around in a series of skips.

Although mudskippers' fins do not have a joint homologous to the elbow, the joint between the radials and the fin rays serves a functionally analogous role.

— Integrative and Comparative Biology^[10]

The mudskipper pectoral fin differs from most actinopterygian fishes in that the radials of the mudskipper pectoral fin are elongate and protrude from the body wall. This unusual morphology creates a pectoral fin with two fin segments (the radials and the rays) and two movable hinge joints: a 'shoulder' joint where the cleithrum meets the radials and an 'intra-fin' joint where the radials meet the rays.^[4] In addition, ... the abductor superficialis muscle of the pectoral fin is divided into two sections (rather than being a single muscle, as is common with the rest of the Oxudercinae gobies) with one section inserting on the dorsal rays and the other section inserting on the ventral rays.

— The Journal of Experimental Biology^[11]

Mudskippers have the ability to breathe through their skin and the lining of their mouth (the mucosa) and throat (the pharynx); this is only possible when the mudskippers are wet, limiting them to humid habitats and requiring that they keep themselves moist. This mode of breathing, similar to that employed by amphibians, is known as cutaneous air breathing.^[5] Another important adaptation that aids breathing while out of water is their enlarged gill chambers, where they retain a bubble of air. These chambers close tightly when the fish is above water, due to a ventromedial valve of the gill slit, keeping the gills moist, and allowing them to function while exposed to air. Gill filaments are stiff and do not coalesce when out of water.^[5]

The different species have adapted to various diets on the mudflats. Boleophthalmus boddarti is detritivorous, while others will eat small crabs, insects, snails and even other mudskippers.^[12]

Digging deep burrows in soft sediments allows the fish to thermoregulate,^[13] avoid marine predators during the high tide when the fish and burrow are submerged,^[14] and lay their eggs.^[15] When the burrow is submerged, several mudskipper species maintain an air pocket inside it, which allows them to breathe in conditions of very low oxygen concentration.^[16][17][18]

Species

Mudskipper at Point Calimere Sanctuary, Tamil Nadu, India

Mudskipper at Coringa Mangrove Sanctuary, Andhra Pradesh, India

Play media

Periophthalmodon septemradiatus territorial defense call and jumping ability^[19]

The genus Periophthalmus is by far the most diverse and widespread genus of mudskipper. Eighteen species have been described.^[20][21][22]Periophthalmus argentilineatus is one of the most widespread and well-known species. It can be found in mangrove ecosystems and mudflats of East Africa and Madagascar east through the Sundarbans of Bengal, Southeast Asia to Northern Australia, southeast China, and southern Japan, to Samoa and Tonga Islands.^[6] It grows to a length of about 9.5 cm ^[6] and is a carnivorous opportunist feeder. It feeds on small prey such as small crabs and other arthropods.^[23] However, a recent molecular study suggests that P. argentilineatus is in fact a complex of species, with at least three separate lineages, one in East Africa, and two in the Indo-Malayan region.^[24]
Another species, Periophthalmus barbarus, is the only oxudercine goby that inhabits the coastal areas of western Africa.^[6]

References

External links

• FishBase entry on Gobiidae


• Gobioid Research Institute


• 
  The Mudskipper: a website on mudskippers or The Mudskipper
  


• Mudskipper photos by Daniel Trim