Gobioidei

(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

List of Species

Large-scale spinycheek sleeper

Eleotris amblyopsis

family

Eleotridae

taxonomy

Culius amblyopsis Cope, 1871, Suriname.

other common names

English: Sleeper.

physical characteristics

Reaches about 3.9 in (10 cm) total length over most of range, largest individual observed from North America about 5.5 in (14 cm). Two dorsal fins, pelvic fins separated, and rounded caudal fin. Torpedo-like body shape with flattened head. Tan to dark brown, sometimes changes to dark trunk laterally with light tan dorsum, especially in juveniles. May have rows of small dark spots on sides. Two dark streaks radiating rearward from eyes on cheeks. A large dark spot on the upper pectoral fin base and some specimens with two elongate spots extending from pectoral fin base onto pectoral fin rays. Stout spine on lower cheek faces forward.

distribution

Common from North Carolina to French Guiana, and in Cuba and Hispaniola. Less frequently known from Brazil and the Antilles.

habitat

Euryhaline (able to live in waters of a wide range of salinities); adults and juveniles inhabit continental estuaries, but are also known from fresh water in Central America and occasionally in the Antilles. Associates with floating, emergent, and marginal vegetation, such as mangroves, river cane, and water hyacinths. Pelagic marine larval stage.

behavior

A lethargic "sit-and-wait" predator. Postlarvae join other species to form large schools which periodically move into estuaries and fresh waters after a marine larval period.

feeding ecology and diet

Predaceous; feeds on shrimps and small fishes.

reproductive biology

Presumably amphidromous, with adults living and reproducing in fresh water or estuaries and a marine larval period.

conservation status

Considered vulnerable in North Carolina at the edge of its range, but not listed by the IUCN.

significance to humans

None known.

Blind cave gudgeon

Milyeringa veritas

family

Eleotridae

taxonomy

Milyeringa veritas Whitley, 1945, Milyering, Yardie, 20 mi (32 km) southwest of Vlamingh Head, North-West Cape, Western Australia. Populations show a low degree of heterozygosity, which is typical of cave-dwelling fishes. This species is possibly closely related to the Indo-Pacific eleotrid genus Butis.

other common names

English: Cave gudgeon.

physical characteristics

Reaches about 2 in (5 cm) total length. The head is broad and concave over the snout. The eyes are not visible. Small first dorsal fin with four very short spines. Second dorsal fin with nine longer rays. Pelvic fins separate. The caudal fin is slightly pointed. The head is naked but the body is covered with small scales (about 28 in series along the flanks). The fish is very poorly pigmented. The top of the head is pale yellow, with a dark gray triangular mark over the brain. The posterior nostrils are pink. There is a purplish spot over the operculum. The body is pale gray and fins are flesh colored.

distribution

Western and northeastern coastal plains of the Cape Range Peninsula and Barrow Island, Western Australia.

habitat

This is a troglobitic species; i.e., an obligatory cave dweller, found in shallow wells (at depths of up to 1.6 ft/0.5m), sink-holes,

and deeper cave systems (at depths of up to 105 ft/32 m) that are freshwater or anchialine (located near the sea and flooded with sea water). In anchialine systems the fishes may live in freshwater layers that overlie the salt water. They are also found in brackish water, tolerating salinities from 0.27–34 parts per thousand and temperatures of 70–83°F (21–28°C).

behavior

Swims slowly, with the pectorals pointed out to the sides and the pelvics extended slightly forward.

feeding ecology and diet

Possibly feeds on insects or other small animals that drop into the cave waters.

reproductive biology

Random mating apparently occurs between subpopulations throughout the range.

conservation status

Listed as Data Deficient in the IUCN 2002 Red List, but as threatened by the Western Australian Conservation Act. Major threats come from habitat disturbance due to urban and agricultural development, and water extraction causing a lowering of the water table. Water quality in the underground systems is being compromised by pollution from nutrients and other chemicals.

significance to humans

None known.

Marble sleeper

Oxyeleotris marmorata

family

Eleotridae

taxonomy

Eleotris marmorata Bleeker, 1852, Bandjarmasin, Borneo, and Palembang, southeast Sumatra, Indonesia.

other common names

English: Marble goby, marbled sand goby; German: Marmorgrundel; Cantonese: Soon hock; Japanese: Kawaanago; Khmer: Trey damrei; Laotian: Pa bou; Malay: Bakutut, belantuk; Tagalog: Bia; Thai: Pla bu jak; Vietnamese: Cá bong.

physical characteristics

The largest gobioid fish, this species may reach 35.4 in (90 cm) total length. Torpedo-like body shape with flattened head. Oblique, terminal mouth. Two dorsal fins and rounded caudal fin. Pelvic fins separate. Body brown with dark mottling giving a marbled appearance.

distribution

Southeast Asia in Laos, Cambodia, Vietnam, Thailand, Malay Peninsula, Sumatra and western Borneo. Also reported from Luzon (Philippines), and introduced into Taiwan for aquaculture.

habitat

Found in rivers, lakes, swamps, ditches, and ponds; water may be muddy or clear and over muddy, sandy, or gravel bottoms. This species may also be found in brackish waters around the mouths of rivers and canals.

behavior

Solitary, nocturnal hunter that prowls slow-moving streams, lakes, and swamps. During the daytime it rests at the bottom, taking cover among rocks, woody debris, or vegetation. This may be the typical behavior for most sleepers, hence the common name.

feeding ecology and diet

Predaceous; primarily eats small fishes, but also takes crustaceans, insects, and mollusks. Larval fish in culture ponds feed on cladocerans, rotifers, chironomids, and brachiopods.

reproductive biology

Reach sexual maturity at approximately 3.9–4.7 in (10–12 cm) standard length. Spawns January through October. The eggs are about 0.09 in (2.3 mm) long and 0.02 in (0.63 mm) wide. Hatching starts at about 41 hours after fertilization, reaching its peak about 60–70 hours after fertilization. The males care for the eggs and guard the newly hatched fry. The larvae are initially pelagic but change to a more benthic habitat after about 25–30 days, becoming quite sedentary after about 40 days.

conservation status

Not listed by the IUCN, but the species may be declining in some parts of its native range due to overharvesting.

significance to humans

A highly prized food fish in Southeast Asia, where it is also raised in ponds. In 2000, 277 tons (282 t) of marble sleeper were produced by aquaculture in Malaysia, Singapore, and Thailand. This quantity represents about 74% of the total global aquaculture of gobioid fishes for 2000, and had a value of about $1.2 million. In Borneo, marble sleepers captured for markets in Singapore and Japan may bring a fisherman $20 per lb ($10 per kg).

Gorgeous prawn-goby

Amblyeleotris wheeleri

family

Gobiidae

taxonomy

Cryptocentrus wheeleri Polunin and Lubbock, 1977, Aldabra Atoll, Seychelles.

other common names

English: Gorgeous goby, Wheeler's shrimp goby; Afrikaans: Mooidikkop; Gela: Iga taotao; Tagalog: Bia.

physical characteristics

Reaches about 3.2 in (8 cm) total length. Body with scales (62–68 in longitudinal series along the flanks). Second dorsal and anal fins, each with 12 soft fin rays. Pelvic fins not forming a strong sucking disk; fins connected between innermost rays but not between spines of each fin. Head and body a light yellowish green or brownish dorsally, with six reddish, vertical, or oblique bands on flanks; numerous small, bluish spots, or reddish spots on head. A vertical red stripe runs from eye to corner of mouth. Fins transparent, and all except pectorals with numerous blue spots and often with reddish bases, and dorsal fins with orange spots near edge. Caudal fin has oblique red stripe.

distribution

Throughout much of the Indo-Pacific region, from East and South Africa and the southern Red Sea to Fiji; in the Pacific, north to Taiwan, and the Ryukyu Islands of southern Japan, and south to the Great Barrier Reef

habitat

Lives on sand and rubble patches of reef flats between 16 and 98 ft (5–30 m) deep. Found in association with alpheid prawns (usually Alpheus ochrostriatus) that live in burrows.

behavior

Typical of several species of gobies that live symbiotically with prawns. The goby rests just outside the prawn's burrow, and retreats into the burrow at the approach of danger. The goby and prawn may stay in reasonably close contact with each other when either leaves the burrow. The goby benefits from the shelter afforded by the burrow, and might also be "cleaned" by the prawn. The goby acts as a "watchman" for the prawn. The goby can signal the presence or absence of danger by flicking movements of its tail, and the prawn feels these signals through its antennae, which it places on the goby's tail. If the prawn cleans the goby of small particles attached to its body, then this may provide a useful food source for the prawn.

feeding ecology and diet

Nothing known.

reproductive biology

Nothing known.

conservation status

Not threatened.

significance to humans

Commercial importance in the aquarium trade.

Whip coral goby

Bryaninops yongei

family

Gobiidae

taxonomy

Cottogobius yongei Davis and Cohen, 1969, Darvel Bay, west of Tatagan Island, Borneo.

other common names

Japanese: Garasuhaze; Samoan: Mano'o.

physical characteristics

Small, reaches about 1.2 in (3 cm) standard length. The preoperculum has a slightly scalloped edge, visible just behind the mouth. Body with scales (26–58 in longitudinal series along the flanks). The pelvics form a small suctorial disk. The head is transparent except for some brownish patches on the sides, a distinct reddish gold stripe that extends dorsally around the snout, and some reddish spots and a band over the top of the head. The eyes are bright golden with reddish rims. The body is transparent, with about six darkish or reddish brown bands on flanks. The ventral part of the body is mottled brown, and there is a large brown or reddish patch at the base of the pectoral fin. The dorsal, anal, and caudal fins may be reddish.

distribution

From the Red Sea, to the Seychelles in the Indian Ocean. More common in the Pacific Ocean from Indonesia east to French Polynesia, north to Hawaii and southern Japan, and south to the Great Barrier Reef.

habitat

Apparently restricted to small patches that are widely dispersed. The fish live only on the sea whip (Cirripathes anguina), found on drop offs in strong currents and on sheltered back reefs, from 9.9 to 148 ft (3–45 m) deep.

behavior

Small sea whips are occupied by a single immature fish. Larger sea whips are occupied by a male-female pair. Juveniles and a second small female may also be present.

feeding ecology and diet

Nothing known.

reproductive biology

The male-female pair on the sea whip are probably monogamous. About 6–8 in (15–20 cm) in from the tip of the sea whip, the fish clear living tissue from the whip in a narrow band about 0.9 in (23 mm) wide, exposing the underlying skeleton. The eggs are attached in a wide band over this "nest." Individual fish can change sex, apparently either way. The largest female to settle on a whip might become a male and the next largest fish remain a female, thus forming a pair. Fish that lose their mates might revert to a bisexual state, allowing them to mate with any fish of either sex that subsequently settles on the sea whip.

conservation status

Not threatened.

significance to humans

Found in the aquarium trade, but not common.

Violet goby

Gobioides broussoneti

family

Gobiidae

taxonomy

Gobioides broussonnetii Lacepede, 1800, Suriname.

other common names

German: Lila Aalgrundel; Spanish: Lamprea; Portuguese: Aimore.

physical characteristics

To about 21.6 in (55 cm) total length. The body is elongate and eel-like. Dorsal fins united to form a long continuous fin confluent with the caudal fin. Anal fin also connected to caudal fin. The caudal fin is long and pointed. Pelvic fins are united to form a concave disk. The mouth is large and oblique. The eyes are small and high on the head. The body and head are purple brown, the abdomen and underside of the head are white. A series of dark chevrons, angled forward, extends along the sides.

distribution

Western Atlantic, Gulf and Caribbean coasts of the Americas, extending from Brunswick County, North Carolina, to Rio Grande do Sul, Brazil. Also on coasts of Cuba, Hispaniola, and Puerto Rico in the Caribbean.

habitat

Lives on muddy bottoms and in burrows; inhabits tidal freshwater and low salinity estuaries offshore from large rivers; one individual was found 298.5 ft (91 m) off the Mississippi River in Louisiana.

behavior

No published information is available, but some is forthcoming from the aquarium trade. Males display for females before spawning by raising and lowering the dorsal fin, and swimming in and around the potential nest site. Males become territorial after spawning and guard the nest.

feeding ecology and diet

The long coiled gut suggests that it feeds on diatoms, algae, and tiny invertebrates sifted from sediments.

reproductive biology

In the aquarium, spawning may last 12–24 hours. Eggs are deposited in nests under an object, presumably within a burrow in the wild. Eggs hatch in 36–38 hours. Males usually guard the fry until they swim freely.

conservation status

Not threatened.

significance to humans

No commercial fishing values, but becoming an increasing part of the aquarium trade because of its interesting appearance.

Neon goby

Gobiosoma oceanops

family

Gobiidae

taxonomy

Elacatinus oceanops Jordan, 1904, Garden Key, Tortugas, Florida.

other common names

English: Blue neon goby.

physical characteristics

Small, slender goby with an pointed, bulbous snout. Maximum length about 2.4 in (6 cm). Has two dorsal fins and a squareshaped caudal fin about the same height as the body. The pelvic fins are joined to form a cuplike sucking disk. Dorsally the fish is dark, with an iridescent pale blue stripe running laterally along the lower edge of the dorsum, extending from the snout to the caudal fin. This lateral streak in fishes from Belize is blue in the middle bordered by white. The underside of the head and the abdomen are pale.

distribution

Southern Florida, the Flower Garden coral reefs 100 mi (161 km) off the Texas-Louisiana border, Alacran Reef of northern Yucatán, Quintana Roo state (Yucatán peninsula of Mexico), and Belize.

habitat

Inhabits coral reefs and tropical rocky substrates at depths from 3.3 to 131 ft (1–40 m). Has also been observed associated with large sponges.

behavior

Rests on coral and withdraws into crevices when threatened. A cleaner goby that picks parasites off other fishes, the male cleans the undersides of rocks, corals, or shells to prepare the surface for eggs.

feeding ecology and diet

This is a cleaner goby that picks parasites off other fishes. As with other cleaner gobies, this species waits for other fishes at stations on top of coral heads. It will then swim along the fish and remove ectoparasites from its skin. Its diet includes parasites, fish scales (perhaps accidentally removed), and benthic invertebrates. Larvae feed on small plankton.

reproductive biology

Males clean the undersides of rocks, corals, or shells to prepare the surface for eggs. The male swims in front of the female and entices her into the shell or crevice. Spawning occurs with the male and female quivering side-by-side, depositing the eggs on the ceiling of the nest. Individuals can spawn multiple clutches, several times within a month. The eggs are about 0.08 in (2 mm) long, and 0.04 in (1 mm) wide. About 300–450 eggs are deposited in a nest about 0.5 sq in (3 sq cm). The pair remains together while caring for the eggs. The male guards the nest and presumably circulates oxygen-rich water over the eggs, using his pectoral and caudal fins. Eggs hatch in 7–10 days. Larvae are 0.16 in (4 mm) at hatching, and the parents do not care for the fry.

conservation status

Not threatened.

significance to humans

This colorful fish is a popular, nonaggressive aquarium species that does not require much space. It is commercially important in the saltwater aquarium trade.

O'opo alamo'o

Lentipes concolor

family

Gobiidae

taxonomy

Sicyogaster concolor Gill, 1860, Hilo, Hawaii. Part of the subfamily Sicydiinae.

other common names

Hawaiian: 'alamo'o.

physical characteristics

To 5.3 in (13.4 cm) total length. A tubular species with a blunt head, subterminal mouth, and rounded caudal fin. Pelvic fins are united to form a cup and adhere to the body. Gray or brownish trunk with irregular lateral markings that sometimes form bands. Breeding males transform from the drab color to black on the anterior half of the body, and red or white tinged with red posteriorly, with white median fins.

distribution

Hawaiian Islands.

habitat

Adults and juveniles ascend waterfalls to inhabit headwaters and upper reaches of perennial rain-forest streams on the high islands of the Hawaiian archipelago. It has been reported from sites as high as 2,947 ft (900 m) above sea level. This species has planktonic marine larvae.

behavior

Larvae of this amphidromous species wash out to sea where they may remain for a period of three months. Post-larvae return to fresh water, and ascend waterfalls to colonize the upper reaches of streams; sicydiine gobies probably use their pelvic fins and their teeth to gain purchase on the rock surface as they climb waterfalls. Genetic evidence suggests that they do not return to the natal stream.

feeding ecology and diet

Predominantly herbivorous, has rows of fine teeth that are ideal for scraping algae from the surface of rocks in the stream bed. As the old teeth break or wear down, they are replaced by fresh teeth. Freshwater shrimps are also taken in small numbers.

reproductive biology

Breeds from October through June. Males may establish and patrol reproductive territories, but evidence for this is inconclusive. Males defend nest cavities prior to breeding and pair with females prior to spawning for two to six days. Spawning may last several hours, during which females deposit eggs on the underside of the roof of the nest cavity before being chased out by the male. Individual females may spawn four clutches a season, with an average of nearly 14,000 eggs. After spawning, males apparently guard the nests. Although not directly observed, males may also clear detritus from nest cavities. Eggs hatch in three to six days.

conservation status

Not currently threatened; classified as Data Deficient by the IUCN. This species was listed as a Category 1 candidate for Endangered Species classification by the U.S. Fish and Wildlife Service from 1989 to 1996. It was removed from the list when it was discovered to be more widely abundant than previously believed. Many populations were not identified prior to listing, in part because upstream sites were undersampled.

significance to humans

No commercial value.

Arno goby

Padogobius nigricans

family

Gobiidae

taxonomy

Gobius fluviatilis var. nigricans Canestrini, 1867, Arno River, Italy.

other common names

Italian: Ghiozzo di ruscello.

physical characteristics

Males reach slightly beyond 4.9 in (12.5 cm) total length. Tubular species with a blunt head and terminal mouth. Two dorsal fins, second with 12–14 fin rays. Rounded caudal fin. Pelvic fins form a sucking disc, which does not extend as far as the anus. There are 44–49 ctenoid scales along the midline of the flanks. Brownish or light brown dorsally, and whitish ventrally. Five characteristic brown bands over the back, the first at the base of the pectoral fin and the last at the caudal peduncle. Dorsal fins with alternating light and dark gray bands; the first dorsal fin has a yellowish orange border.

distribution

Formerly widespread through fresh waters of central Italy. Currently found in the Serchio, Arno, Ombrone, and Tiber River basins. Populations have become restricted because of environmental degradation and pollution.

habitat

Small to medium-sized streams with moderate current. Usually found in clear waters, over a stony substrate.

behavior

Usually stationary, positioned under stones, occasionally moves quickly from one stone to another.

feeding ecology and diet

Predaceous; feeds on oligochaetes, crustaceans, and insect larvae.

reproductive biology

Probably reaches sexual maturity during the first year, at 1.6–2 in (4–5 cm). Males become very dark during the reproductive period, which usually occurs between May and June. The male is territorial and guards a nesting cavity under a stone, attracting females with a courtship display and making noises as he does. The male carries small stones out from the nest in his mouth and deposits them near the female, and then quickly returns to the nest. He repeats this behavior until the female enters the nest, and then swims several times upside down over the roof of the nest. The female attaches herself to the roof of the nest using the pelvic disk and spawns 100–200 eggs, which are stuck to the roof. Several females can spawn in the same nest, which may hold over 1,500 eggs at various stages of development. The male cares for the eggs until they hatch, after about 17 days.

conservation status

Listed as Vulnerable by the IUCN, with a suspected population reduction of at least 20% over the last 10 years, or three generations. Populations are threatened by environmental degradation and, in some areas, by the introduction of Padogobius martensii, which probably competes for similar ecological niches.

significance to humans

There is no commercial or sports fishery for this species. However, the flesh tastes good and fishes are collected illegally and eaten fried.

Dwarf pygmy goby

Pandaka pygmaea

family

Gobiidae

taxonomy

Pandaka pygmaea Herre, 1927, Philippines (possibly Malabon).

other common names

German: Zwerggrundel; Tagalog: Bia.

physical characteristics

A very small fish; females reach 0.6 in (1.5 cm) total length. Body is covered with scales, up to 25 in a longitudinal series along the flanks. The second dorsal and anal fins have six or seven soft rays. The pelvic fins are united into a small sucking disk. The body has a conspicuous pigmentation, with about four cross bands of dark spots on the flanks, and heavy pigmentation on the bases of all the fins except the pelvics. In males, the first dorsal fin is tipped with blue and the lower third is yellowish.

distribution

First reported as possibly from Malabon, a suburb of Manila in the Philippines. The presence of the species in the Malabon River was confirmed in the 1950s. Subsequently, it has been found at an island off Palawan (Philippines), the east coast of peninsular Thailand, Singapore, Bali, Sulawesi, and New Guinea.

habitat

Found along river banks in the Philippines from January to June, when the water is clear and the tide is rising. During the rainy season the fishes apparently move into ponds and canals. Also found in brackish waters and around mangroves. Some specimens were apparently caught in marine waters adjacent to Palawan. Fishes kept in aquaria can acclimate to marine, brackish, or freshwater conditions.

behavior

Usually stationary, on surfaces of rocks or plants, or hanging in midwater. Swimming occurs in an erratic series of starts.

feeding ecology and diet

Aquarium-kept specimens are voracious, feeding on brine shrimps and other small or chopped invertebrate material.

reproductive biology

Nothing known.

conservation status

Listed as Critically Endangered by the IUCN, with an observed or suspected population reduction of 90% or more over the last 10 years, or three generations. Populations are threatened by introduced species, hybridization, pathogens, pollutants, competitors, or parasites.

significance to humans

None known.

Atlantic mudskipper

Periophthalmus barbarus

family

Gobiidae

taxonomy

Gobius barbarus Linnaeus, 1766, Liberia. Based on a neotype specimen designated in 1989.

other common names

French: Sauteur de vase atlantique; German: Schlammspringer; Spanish: Saltafango atlantico; Akan: Soetsi; Portuguese: Saltaoda-vasa.

physical characteristics

To about 6 in (15 cm) total length. Prominent eyes with ventral eyelids perched on top of large head. Two dorsal fins. Muscular pectoral fin and pelvic fin bases used for crawling and climbing. Pelvic fins separate. Body tan but light ventrally, and with black diagonal bars dorsally on flanks; opalescent spots on head and anterior trunk; light margin on first dorsal fin may be tinged with light blue, lower portion dark; second dorsal fin with dark submarginal band over a light band.

distribution

West African coast from Senegal to Angola, including most islands.

habitat

Inhabits the littoral zone of mangrove estuaries and muddy intertidal flats where it lives in burrows.

behavior

Amphibious; lives in burrows in the mangal zone. Territorial, forages for food on the mudflats and around the mangroves themselves. They can flee predators by skipping or hopping across the flats and into the mangrove forests or into their burrows. When on land, mudskippers keep a mouthful of water for extracting oxygen via the gills, and they can breathe through the skin which is well served with blood vessels.

feeding ecology and diet

Eats crustaceans, worms, and insects from the intertidal zone.

reproductive biology

Reproduction occurs in burrows.

conservation status

Not threatened.

significance to humans

No commercial fishing value; plays a minor role in the aquarium trade.

Samoan sand dart

Kraemeria samoensis

family

Kraemeriidae

taxonomy

Kraemeria samoensis Steindachner, 1906, Samoan Islands.

other common names

English: Sand dart; Afrikaans: sandspies.

physical characteristics

Small, elongate fish, reaches 1.4 in (3.5 cm) total length. Lacks scales. Head with minute eyes, and a "chinlike" forward projection of the lower jaw. Six or seven small flaps project from the lower edge of the preoperculum, just behind the mouth, and five or six flaps project from the lower edge of the operculum covering the gills. There is a single, long, dorsal fin, and a long anal fin. Specimens from the Indian Ocean have fewer opercular flaps than those from the Pacific, and might represent a different species.

distribution

The Indo-Pacific region, from East Africa to the Society Islands.

habitat

Buries into loose coral sand in inshore areas where there is strong wave action.

behavior

Nothing known.

feeding ecology and diet

Feeds on polychaetes.

reproductive biology

Nothing known.

conservation status

Not threatened.

significance to humans

None known.

Fire goby

Nemateleotris magnifica

family

Ptereleotridae

taxonomy

Nemateleotris magnificus Fowler, 1938, Buka Buka Island, Gulf of Tomini, Sulawesi, Indonesia.

other common names

English: Fire dartfish, firefish; French: Gobie de feu; Afrikaans: Vuur-dikkop; Japanese: Hatatatehaze; Malay: Roket antene; Samoan: Mano'o-sugale.

physical characteristics

Maximum size 3.5 in (9 cm) total length. This is one of the "hovering" gobies, a species that actually hovers in the water column instead of hugging the bottom like most of its kin. The body form is more compressed laterally (deeper and thinner) instead of the more typical tubular goby shape. The anterior portion of the first dorsal fin is greatly elongate, almost as long as the fish itself in some specimens. The pelvic fins are separate. The second dorsal and anal fins are long and, when extended, have the appearance of a feather, or the tail end of an arrow or a dart. The fish has a rounded caudal fin. The head has a yellow mask over the snout and eyes. The front half of the fish (other than the mask) is white, the back half grades from orange to red, with green streaks in the median fins converging posteriorly.

distribution

Tropical Indian and Pacific Oceans, from South Africa to the Marquesas. In the Pacific, extends north to the Hawaiian and Ryukyu Islands, and south to New Caledonia.

habitat

Coral-reef species found on the upper slopes of the outer reef. Although observed at depths from 20–200 ft (6–61 m), it is usually found at less than 92 ft (28 m). Creates burrows to which it can retreat when threatened.

behavior

Occurring singly or in small group, this species hovers in the water column, a few centimeters above sand or rubble, selectively feeding on drifting zooplankton. Usually has a small territory around a hole, cave, or burrow into which it retreats at the threat of danger. The burrow may be shared. Has a habit of flicking the first dorsal spine up and down, erecting the dorsal fin when threatened or defending territory.

feeding ecology and diet

Feeds on zooplankton, including crustacean larvae and copepods.

reproductive biology

Spawning occurs in burrows.

conservation status

Not threatened.

significance to humans

This beautiful fish is an important saltwater aquarium species.

Loach goby

Rhyacichthys aspro

family

Rhyacichthyidae

taxonomy

Platyptera aspro Valenciennes, 1837, west Java and Sulawesi. Although Rhyacichthys aspro has several specializations of its own, it is considered to be a primitive representative of the Gobioidei.

other common names

Ilokano: Kampa; Japanese: Tsubasahaze; Visayan: Dalapakan.

physical characteristics

Reaches 12.6 in (32 cm) total length. Head depressed dorsally, with a wedgelike snout, and with small eyes. Mouth ventrally placed on head with a fleshy upper lip. Body laterally compressed toward tail, which is slightly forked; body with 27–40 scales in longitudinal series on flanks. Lateral line system well developed on head and body. Pelvic fins separate, each with a patch of enlarged musculature on the anterior border. Body light brown with several darker, longitudinal stripes on flanks; dark bands running from eye toward anterior tip of snout. Dorsal fins with alternating light and dark longitudinal stripes; caudal and pectoral fins pale with several dark vertical bands.

distribution

Islands of the western Pacific rim, extending in an arc from the Ryukyu Islands of southern Japan, through Taiwan, Indonesia, and the Philippines, to New Guinea and the Solomon Islands.

habitat

Clings to rocks and boulders in fresh waters of torrential hill streams. The streamlined head and body shape makes this species well suited to living in these fast-flowing waters. The fish may use the ventral part of head, thoracic region of the body, and the pectoral and pelvic fins to attach to rocks. This might be done by friction between the rock surface and rough surfaces on the underside of the body, or by suction, in which a low-pressure water flow between the body and the rock "sticks" the fish to the substrate.

behavior

Swims in a series of rapid darts and, at the approach of danger, quickly moves into crevices under large rocks or boulders.

feeding ecology and diet

Herbivorous, browses on algae on the surfaces of stones.

reproductive biology

Nothing known.

conservation status

Not threatened.

significance to humans

None known.

Resources

Books

Akihito, Prince. "Some Morphological Characters Considered to Be Important in Gobiid Phylogeny." In Indo-Pacific Fish Biology: Proceedings of the Second International Conference on Indo-Pacific Fishes, edited by T. Uyeno, R. Arai, T. Taniuchi, and K. Matsuura. Tokyo: Ichthyological Society of Japan, 1986.

Akihito, Prince, M. Hayashi, T. Yoshino, K. Shimada, H. Senou, and T. Yamamoto. "Suborder Gobioidea." In The Fishes of the Japanese Archipelago, edited by H. Masuda, K. Amoaka, C. Araga, T. Uyeno, and T. Yoshino. Tokyo: Tokai University Press, 1984.

Colin, P. The Neon Gobies; the Comparative Biology of the Gobies of the Genus Gobiosoma, Subgenus Elacatinus, (Pisces: Gobiidae) in the Tropical Western North Atlantic Ocean. Neptune City, NJ: T. F. H. Publications, 1975.

Harrison, I. J., and M. L. J. Stiassny. "The Quiet Crisis: A Preliminary Listing of the Freshwater Fishes of the World That Are Extinct or 'Missing in Action.'" In Extinctions in Near Time: Causes, Contexts, and Consequences, edited by R. D. E. MacPhee. New York: Kluwer Academic/Plenum Publishers, 1999.

Hoese, D. F. "Gobies." In Encylopedia of Fishes, edited by J. R. Paxton and W. N. Eschmeyer. San Diego: Academic Press, 1995.

——. "Gobioidei: Relationships." In Ontogeny and Systematics of Fishes, edited by H. G. Moser. Gainesville, FL: American Society of Ichthyologists and Herpetologists, 1984.

Miller, P. J. "The Tokology of Gobioid Fishes." In Fish Reproduction: Strategies and Tactics, edited by G.W. Potts and R. J. Wooton. London: Academic Press, 1984.

—— "Reproductive Biology and Systematic Problems in Gobioid Fishes." In Indo-Pacific Fish Biology: Proceedings of the Second International Conference on Indo-Pacific Fishes, edited by T. Uyeno, R. Arai, T. Taniuchi, and K. Matsuura. Tokyo: Ichthyological Society of Japan, 1986.

Myers, R. F. "Suborder Gobioidei." In Micronesian Reef Fishes: A Comprehensive Guide to the Coral Reef Fishes of Micronesia, 3rd edition. Guam: Coral Graphics, 1999.

Nelson, J. S. Fishes of the World, 3rd edition. New York: John Wiley & Sons, 1994.

Nishimoto, R. T., and J. M. Fitzsimons. "Courtship, Territoriality, and Coloration in the Endemic Hawaiian Freshwater Goby, Lentipes concolor." In Indo-Pacific Fish Biology: Proceedings of the Second International Conference on Indo-Pacific Fishes, edited by T. Uyeno, R. Arai, T. Taniuchi, and K. Matsuura. Tokyo: Ichthyological Society of Japan, 1986.

Patterson, C. "Osteichthyes: Teleostei." In The Fossil Record, Vol. 2, edited by M. J. Benton. London: Chapman and Hall, 1993.

Torricelli, P. "Gobiidae." In I pesci delle acque interne italiane, edited by G. Gandolfi, S. Zerunian, P. Torricelli, and A. Marconato. Instituto Poligrafico e Zecca dello Stato, Italy: Ministero dell'Ambiente e Unione Zoologica Italiana, 1991.

Periodicals

Akihito, A. Iwata, T. Kobayashi, K. Ikeo, T. Imanishi, H. Ono, Y. Umehara, C. Hamamatsu, K. Sugiyama, Y. Ikeda, K. Sakamoto, A. Fumihito, S. Ohno, and T. Gojobori. "Evolutionary Aspects of Gobioid Fishes Based upon a Phylogenetic Analysis of Mitochondrial Cytochrome B Genes." Gene 259, nos. 1 and 2 (2000): 5–15.

Allen, G. R., and D. F. Hoese. "A Review of the Genus Mogurnda (Pisces: Eleotrididae) from New Guinea with Descriptions of Three New Species." Ichthyological Exploration of Freshwaters 2, no. 1 (1991): 31–46.

Erdman, D. S. "The Green Stream Goby, Sicydium plumieri, in Puerto Rico." Tropical Fish Hobbyist (Oct. 1986): 70–74.

Hoese, D. F., and A. C. Gill. "Phylogenetic Relationships of Eleotridid Fishes (Perciformes: Gobioidei)." Bulletin of Marine Science 52, no. 1 (1993): 415–440.

Larson, H. K. "A Review of the Australian Endemic Gobiid Fish Genus Chlamydogobius, with a Description of Five New Species." The Beagle, Records of the Museums and Art Galleries of the Northern Territory 12 (1995): 19–51.

——. "A Revision of the Gobiid Genus Bryaninops (Pisces), with a Description of Six New Species." Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 2(1985): 57–93.

Miller, P. J. "The Adaptiveness and Implications of Small Size in Teleosts." Symposia of the Zoological Society of London 44(1979): 263–306.

——. "The Endurance of Endemism: the Mediterranean Freshwater Gobies and Their Prospects for Survival." Journal of Fish Biology 37, supp. A (1990): 145–156.

——. "The Functional Ecology of Small Fish: Some Opportunities and Consequences." Symposia of the Zoological Society of London 69 (1996): 175–199.

——. "Grading of Gobies and Disturbing of Sleepers." NERC News (Oct. 1993): 16–19.

——. "The Osteology and Adaptive Features of Rhyacichthys aspro (Teleostei: Gobioidei) and the Classification of Gobioid Fishes." Journal of Zoology (London) 171 (1973): 397–434.

Munday, P. L., S. J. Pierce, G. P. Jones, and H. K. Larson. "Habitat Use, Social Organization and Reproductive Biology of the Seawhip Goby, Bryaninops yongei." Marine and Freshwater Research 53 (2002): 769–775.

Murdy, E. O. "A Review of the Gobioid Fish Genus Gobioides." Ichthyological Research 45, no. 2 (1998): 121–133.

——. "A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae: Oxudercinae)." Records of the Australian Museum 11 (1989): 1–93.

Norman, A. "The Smallest Fish There Is." Freshwater and Marine Aquarium 4, no. 1 (1981): 26–27, 82.

Polunin, N. V. C., and R. Lubbock. "Prawn-Associated Gobies (Teleostei: Gobiidae) from the Seychelles, Western Indian Ocean: Systematics and Ecology." Journal of Zoology (London) 183 (1977): 63–101.

Shibukawa, K., A. Iwata, and S. Viravong. "Terateleotris, a New Gobioid Fish Genus from Laos (Teleostei, Perciformes), with Comments on Its Relationships." Bulletin of the National Science Museum, Tokyo Series A 27, no. 4 (2001): 229–257.

Thacker, C. "Phylogeny of the Wormfishes (Teleostei: Gobioidei: Microdesmidae)." Copeia 2000, no. 4 (2000): 940–957.

Tyler, J. C. and J. E. Bohlke. "Records of Sponge Dwelling Fishes, Primarily of the Caribbean." Bulletin of Marine Science 22 (1972): 601–642.

Winterbottom, R. "Search for the Gobioid Sister Group (Actinopterygii: Percomorpha)." Bulletin of Marine Science 52, no. 1 (1993): 395–414.

Other

"Taxonomy Browser." National Center for Biotechnology Information Databases [cited January 14, 2003]. <http://www.ncbi.nlm.nih.gov/Taxonomy/taxonomyhome.html>

FishBase [cited January 14, 2003]. <http://www.fishbase.org/search.cfm>

"FISHSTAT Plus: Universal Software for Fishery Statistical Time Series." Food and Agriculture Organization of the United Nations, Fisheries Department, Fishery Information, Data and Statistics Unit [cited January 14,2003]. <http://www.fao.org/fi/statist/FISOFT/FISHPLUS.asp>

Harrison Ian, J., and Melanie L. J. Stiassny. "List of Fish Extinctions Since a. d. 1500." Committee on Recently Extinct Organisms (CREO) [cited January 14, 2003]. <http://creo.amnh.org/pdi.html>

Ian J. Harrison, PhD

Frank Pezold, PhD