Gobioidei (Gobies)

(vertebrate zoology) The gobies, a suborder of morphologically diverse actinopterygian fishes in the order Perciformes; all lack a lateral line.

(Gobies)

Class: Actinopterygii

Order: Perciformes

Suborder: Gobioidei

Number of families: 2–9

Evolution and systematics

The suborder Gobioidei contains about 2,121 species in about 268 genera, representing about 9% of living teleostean species. Several different groups of acanthomorphs (spinyrayed teleosts) have been suggested as the group to which gobioids are most closely related, but the proposed relationships are usually based on relatively little evidence. A morphological study in 1993 suggested that gobioid fishes might be related to one of the following: the gobiesocids (clingfishes) with the callionymoids (dragonets); the trachionoids (weeverfishes and their relatives); or the hoplichthyids (ghost flatheads).

The taxonomy and phylogenetic (evolutionary) relationships within the Gobioidei are also poorly known. It is unsurprising, therefore, that the classification of gobioid fishes has been described as "chaotic." Since 1973 they have been classified in as few as two families, Rhyacichthyidae and Gobiidae; or as many as nine families: Rhyacichthyidae, Odontobutidae, Eleotridae (called Eleotrididae by some authors), Xenisthmidae, Microdesmidae, Ptereleotridae, Kraemeriidae, Schindleriidae, and Gobiidae (including the subfamilies Gobiinae, Amblyopinae, Oxudercinae, Sicydiinae, and Gobionellinae).

Gobioid fishes are characterized by a lack of several bones in the head and axial skeleton, the presence of free sensory neuromasts borne on raised papillae on the head, and sperm-duct glands associated with the testes. The Rhyacichthyidae, or loach gobies, comprising just two species, is accepted as the most primitive gobioid group. The remaining gobioid groups share several features not seen in the Rhyacichthyidae; for example, the modification of the bones in the head, the reduction of the lateral line canal on parts of the head, and the complete loss of the lateral line canal on the body.

Several features of the skeleton and scales unite the seven families of higher gobioids (Eleotridae, Xenisthmidae, Microdesmidae, Ptereleotridae, Kraemeriidae, Schindleriidae, and Gobiidae) into one large group. Some authors apply the name Gobiidae to this whole group, which does not include the Odontobutidae. However, molecular analyses of the cytochrome b genes in gobioid fishes (made in 2000), and morphological analyses (made in 2002) indicate that the Odontobutidae, as distinct from the other higher gobioids, is an artificial assemblage (i.e., the characters that apparently unite species in the Odontobutidae are not genuinely comparable across all the species, or the characters are not restricted only to those species but may also be found in other, unrelated species).

Since 1998 two new gobioid genera, Protogobius and Terateleotris, have been described with lateral line canals extending onto the body (a primitive feature that is otherwise restricted to the Rhyacichthyidae). The affinities of these two genera to other gobioid fishes are problematic, and neither has been formally assigned to a family. Morphological evidence suggests similarities between Protogobius and Rhyacichthys, but this is based mainly on shared primitive features. Analysis of mitochondrial cytochrome b genes also showed that Protogobius and Rhyacichthys appear to be closely related. Terateleotris lacks several of the derived features that are shared by the seven families of higher gobioids, so this group may be related to either of the more basal groups, Rhycichthyidae or Odontobutidae.

A late Cretaceous or early Tertiary origin of gobioid groups has been proposed. Fossil otoliths that might have come from gobioid fishes have been found in the Harudi Formation of Katchchh, India (Lutetian age; 43.6–52 million years old) and the Naggulan Formation of Java (Bartonian age; 40–43.6 million years ago). The oldest reliably identified skeletal remains are of Pomatoschistus (?) cf. bleicheri (family Gobiidae), from the Headon Hill Formation, Isle of Wight, Great Britain, and date to the Priabonian age (36.6–40 million years old). The oldest fossil material of any eleotrid gobies are otoliths from Faluns de Gaas, Landes, France, and date to the Rupelian age (30–36.6 million years old).

Gobioids play a particularly important role in the freshwater fish communities of oceanic islands, as well as long-isolated fragments of Gondwana such as Madagascar and Australia.

Physical characteristics

Gobioids are small to medium size fishes, with a cylindrical or laterally compressed body. Most species are between about 1.6 and 3.9 in (40–100 mm) in size but several species are smaller. Approximately 40% of gobioid genera include species under 1.6 in (40 mm) maximum body length, and 22% include species under 1.2 in (30 mm). Trimmatom nanus (family Gobiidae) is the shortest known vertebrate at sexual maturity, with females ripening at 0.3 in (8 mm). Schindleria praematura (family Schindleriidae) is the lightest known vertebrate at sexual maturity, not weighing more than 0.03 oz (0.8 g). There are some examples of large gobioid fishes, although these are rare and they are usually freshwater species. For example, the fat sleeper (Dormitator maculatus) and the marble sleeper goby (Oxyeleotris marmorata) in the family Eleotridae are reported to reach over 23.6 in (60 cm) total length.

The gobioid head is usually short and broad, and may be depressed (flattened dorsally). The eyes are positioned on the dorsolateral part of the head. There are usually two separate dorsal fins, the first with five or more weak spines (typically six), and the second dorsal fin with one weak spine and five or more soft, segmented rays. The anal fin also usually has one weak spine and five or more soft, segmented rays. The pectoral fins are generally broad and rounded, but may appear pointed in some species. The paired pelvic fins are positioned thorasically, just below the pectoral fins. Each pelvic fin has one spine and five, soft, segmented rays. The caudal fin is usually broad and rounded, but may appear long and lanceolate in some species. Most species have scales covering the body, although some species lack scales or have them restricted to the posterior part of the body. The size of the scales varies between species, from small and sometimes well embedded in the skin to relatively large. Coloration is variable between species; some are very brightly colored, whereas many others are shades of brown or off white. Both sexes possess a well-differentiated urogenital papilla, posterior to the anus. The urogenital papilla is short and rounded in mature female fish, whereas it is longer and more pointed in mature males.

Gobioid fishes have free, sensory neuromast organs that are distributed on the head in rows of raised papillae. The rows are arranged in diagnostic patterns that are variations of two basic types; a longitudinal arrangement (where all the rows on the cheek run horizontally along the cheek); and a transverse arrangement (where several of the rows run vertically down the cheek). Precise similarities in the arrangement of the papillae rows have been used to infer phylogenetic relationships between groups and hence to classify species. Many gobioid fishes have the pelvic fins united to form a cupshaped, weakly suctorial disc; this is typical of most, but not all of the Gobiidae, and some of the Microdesmidae and Kraemeriidae. The pelvic fins are separate in the Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotrididae, and Ptereleotridae, and are absent in Schindleriidae.

Distribution

Gobioid fishes are common in the tropics, but may also be found in subtropical and temperate regions. They are absent from polar regions. Most of the species are found in marine coastal waters and in estuaries. They are usually in shallow waters, although some species have been found at depths of about 2,625 ft (800 m). About one-tenth of the known species inhabit fresh waters, including the Orinoco and Amazon River basins in South America, and the Nile, Niger, and Congo River basins in Africa. Some species have very widespread distributions. For example, the dusky sleeper (Eleotris fusca) is found in fresh, brackish, and marine waters of the Indo-Pacific, from East Africa through to French Polynesia in the South Pacific. In contrast, some other species have very restricted distributions.

Habitat

Gobioid fishes are found in several diverse marine and brackish-water habitats. About half the known species of gobioid fishes are found in coral reef habitats. Gobiid genera such as Gobiodon and Tenacigobius spend much of their life among the branches of corals such as the black or thorny corals (Antipatharia) and the sea fans and sea whips (Gorgonacea). Gobiosomine gobies (family Gobiidae) such as Evermannichthys, Risor, and Gobiosoma, include tiny species that live exclusively within sponges.

Many gobiid species live epibenthically (on the surface) over sandy or muddy substrates. European species of Gobius and Pomatoschistus are very common in these habitats. Mudskippers (for example, some species of the gobiid genus Periophthalmus) are benthic inhabitants of mangroves. They are highly amphibious, crawling out onto the mud or onto the mangrove roots, and some species also construct burrows, into which they retreat at high tide. Representatives of several other groups of gobioid fishes exhibit a burrowing lifestyle. For example, species of Kraemeriidae (or sand gobies) and the eleotrid genus Calumia burrow in sand; the gobiid Luciogobius burrows in gravel, and species of amblyopine gobiids burrow in mud. Some species, such as the blind goby (Typhlogobius californiensis), from the Eastern Pacific, and the arrow goby (Clevelandia ios), live inside the burrows of marine invertebrates.

There are a few examples of blind eleotrid and gobiid species that live in the subterranean fresh waters of caves and sinkholes; for example, Oxyeleotris caeca from Papua New Guinea, Glossogobius ankaranensis from Madagascar, and the blind cave gudgeon (Milyeringa veritas) from Australia. The two species of loach goby (Rhyacichthys aspro and Rhyacichthys guilberti) show several anatomical specializations for life in torrential rivers and streams. Although the Rhyacichthyidae represents the most basal gobioid lineage, this is not taken as evidence that ancestral gobies also lived in fast-flowing rivers. Instead, it is assumed that this habitat is a specialization of the Rhyacichthyidae. Many species of the Sicydiinae (a sub-family of Gobiidae), including o'opo alamo'o (Lentipes concolor), also inhabit torrential hill streams. Their pelvic fins form a strong suctorial disc that allows them to attach firmly to rocks, preventing the fish from being swept away by the swift currents. Some gobies, such as species of Chlamydogobius, live in freshwater desert habitats. The Dalhousie goby (Chlamydogobius gloveri) can tolerate temperatures up to 111°F (43.9°C); the desert goby (Chlamydogobius eremius) can tolerate temperatures between 41 and 106°F (5–41°C), salinities in the range 0–60 parts per thousands and oxygen levels as low as 0.8 parts per million.

Some gobiid species are nektonic, living in the midwater region of shallow coastal waters and estuaries. Cool-temperate midwater dwellers include small aphyiine gobies (family Gobiidae) such as the transparent goby (Aphia minuta) and the crystal goby (Crystallogobius linearis). Some small nektonic species are also found in warm-temperate and tropical regions; for example, species of Parioglossus are found around mangrove roots.

Behavior

Some nektonic species form schools. For example, species of Parioglossus or Gobiopterus may form small schools around mangrove roots. Epibenthic species are less likely to form schools; however, large populations (in terms of number of individuals) of epibenthic species may be found in some ecosystems. For example, the common goby is densely distributed through some English estuaries. Some species, such as sicydiines, have a larval phase that migrates upriver in schools of very large numbers. The adults of many species are territorial, especially during the breeding season. Some species live in a commensal relationship with invertebrate species. For example, the gorgeous prawn goby (Amblyeleotris wheeleri) lives in the burrows of alpheid shrimps. Other species, such as the neon goby, act as "cleaner" fish, picking parasites off large fishes. Mudskippers (e.g., Periophthalmus barbarus) are best known for their amphibious lifestyle.

Feeding ecology and diet

Gobioid fishes commonly feed on invertebrates. Epibenthic species feed on crustaceans, small worms, and insect larvae associated with the benthos. Some coral-dwelling gobies, such as gobiodontines, feed on the polyps of the corals on which they live. Nektonic species living in the water column, such as the glass goby (Gobiopterus chuno) from fresh and brackish waters of Asia, feed on plankton. Large eleotrid gobies may feed on other, smaller fish. Various gobioid fish species are specialized for grazing on algae. Sicydiine gobies, such as oopo alamoo have rows of fine teeth that are ideal for scraping algae off rock surfaces. Some algal feeders have elongate, elaborately coiled guts to assist in the digestion of the tough algal material, e.g., species of Chlamydogobius from Australia and Kelloggella from the Western Pacific. "Cleaner" gobies, such as the seven-spine, gobiosomine gobies (e.g., Gobiosoma genie and the neon goby Elacatinus oceanops) are specialized feeders that pick parasites off the scales and skin of other fish, sometimes entering the mouth and gills to extract food items. Another seven-spine goby, the nineline goby (Ginsburgellus novemlineatus) shelters on rocks beneath the test, or outer skeleton, of the sea urchin (Echinometra locunter) and feeds on the tube feet of the urchin.

The small size of many gobioid species allows them to exploit meiobenthic food resources that are too small for other species. The small size of the gobies might also put them below the threshold for attack by some large, predatory fishes. However, the small size also places the fish at risk from a variety of other predators. For example, coral-dwelling gobiid species of Eviota less than 1.2 in (30 mm) long may be eaten by slightly larger coral-dwelling cardinalfishes 1.8–2.1 in (45–53 mm) long. The nineline goby, which hides under sea urchins, can fall prey to long-snouted predators like trumpetfishes, that can delve between the spines of the urchin. Large invertebrates, such as shrimps, can overpower small fishes, and there is a reported case of a small goby being ingested by the nemertean worm Lineus longissimus. Small reef-dwelling gobies regularly fall prey to piscivorous mollusks of the genus Conus. Freshwater gobioids are regularly preyed upon by water snakes and fish-eating birds.

Reproductive biology

The male may defend a small territory and nest site that, frequently, is a small space under some stones. Often the male is responsible for caring for the developing eggs. The female will lay a few to several hundred small eggs, attaching them to the underside of rocks, vegetation, or onto corals. The eggs usually hatch in a few days, and the young may be dispersed by water currents. In freshwater species such as sicydiine gobies (e.g., o'opo alamo'o), the larvae are probably swept downstream by the water current; they spend a few weeks to months at sea before returning to fresh water. Frequency of spawning is very variable between different species of gobies. Some species, such as the Japanese ice goby (Leucopsarion petersi), are typically semelparous species, spawning once in a single spawning season, before dying. Other species are iteroparous, relying on more than one spawning event during their lifespan to achieve a satisfactory reproductive output. This "repeat-spawning" behavior is often seen in small species that face reasonably high predation and a short lifespan. The common goby (Pomatoschistus microps), which grows up to 3.5 in (9 cm) total length and can convert 40% of its food into gonad energy, may spawn up to six times over a three-month period, or as many as nine times during its 18-month life cycle. In contrast, the burrowing species Fries' goby (Lesuerigobius friesii) reaches only 1.6 in (4 cm) longer than P. microps, but it does not mature until it is two years old, spawns just twice per season, and may live for 11 years. Egg size and number can also vary widely between species of gobies. The smallest eggs are 0.01 in (0.26 mm) in diameter, produced by the empire gudgeon (Hypseleotris compressa) from Australia and New Guinea; the largest are 0.1 in (2.4 mm) in diameter produced by the knout goby (Mesogobius batrachocephalus) from the Black and Caspian Seas. Some small gobies that live in or around corals and sponges may change sex, which ensures that there are always enough individuals of both sexes to ensure reproduction and survival of the population.

Conservation status

The 2002 IUCN Red List of Threatened Species includes five gobioid species ranked as Critically Endangered and 26 ranked as Vulnerable. Another 21 species are ranked under Lower Risk/Conservation Dependent or Lower Risk/Near Threatened, and 27 are listed as Data Deficient (i.e., there is inadequate information to make a direct, or indirect, assessment of the risk of extinction).

The critically endangered species are threatened by the degradation of their habitat and the introduction of invasive species. For example, Chlamydogobius micropterus is from small springs, bores, and drains in Queensland, Australia, that are threatened by water extraction. The Edgbaston goby (Chlamydogobius squamigenus) is from similar Queensland habitats that are being degraded by trampling from humans and domestic livestock. The introduction of the mosquitofish (Gambusia holbrooki) presents another threat to C. squamigenus. The poso bungu (Weberogobious amadi), which is restricted to Lake Poso in Sulawesi, was probably last observed in 1985 and might be extinct. It has been threatened by pollution, the introduction of alien fish species to the lake and by diseases carried by those introduced species. The sinarapan (Mistichthys luzonensis) from the Philippines was previously thought to be possibly extinct because of the introduction of tilapia to Philippine fresh waters. However, it is still found in lakes of Luzon and is ranked as Lower Risk/Conservation Dependent by the IUCN.

The 26 species listed as Vulnerable include several that have restricted ranges. For example, four of the eight eleotrid species listed as Vulnerable are species of Mogurnda that are restricted to a single lake (Lake Kutubu) in Papua New Guinea. The gobiid Economidichthys trichonis, which is endemic to Lake Trichonis in Greece, is threatened by agricultural activities that may cause eutrophication of the lake. Knipowitschia punctatissima from Italy is threatened by industrial and agricultural activity, and Knipowitschia thessala from Greece is threatened because some of its native watercourses are now dry.

Significance to humans

Gobioid fishes do not support important commercial fisheries, but subsistence fisheries exist for some species in the tropical Atlantic and Indo-Pacific. The sinarapan is a delicacy around the Philippine lakes where it is found, and fry of sicydiine gobies are a delicacy in parts of the Caribbean and Indo-Pacific, where they may be made into a paste. According to FAO statistics, the 2000 global capture for all gobioid fishes was 51,199 tons (52,021 tonnes); 63% of this was caught in the northwest Pacific by countries of the Russian Federation, and 15% was caught in the western Central Pacific by the Philippines. Aquaculture produced 376 tons (382 tonnes) of gobioid fishes in 2000, with a value of $3.38 million. Many gobioid species probably represent important food resources to larger fish species that are commercially fished.

Gobioid fishes form a small part of the ornamental fish trade which, by 1992, had a global retail trade value of $3,000 million. In Hong Kong, a major distributor of ornamental fishes, gobies represented only about 1% of the total number of fishes observed in a market survey in 1996 and 1997. Gobies are an important complement of the biological diversity of coral reefs, which are popular sites for tourism. Therefore, these reef-dwelling gobies represent an indirect source of sustainable income to tropical countries that are using their reefs as attractions for properly managed ecotourism.

Introduced species of gobies include the round goby (Neogobius melanostomus) in the Great Lakes region of North America; the Japanese goby (Tridentiger trigonocephalus), introduced to North American and Australian coastal waters; and the Indo-Pacific Butis koilomatodon, introduced into coastal waters of Nigeria, Panama, and Brazil. Translocated Glossogobius giuris and Hypseleotris agilis appear to have played a role in the extirpation of the endemic cyprinid species flock of Lake Lanao in the Philippines. Native fish species may suffer from competition with these alien species, and the economic cost of monitoring the spread of exotics, and attempting to control them, may be significant.

Species accounts

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[Article by: Ian J. Harrison, PhD; Frank Pezold, PhD]