freshwater fish distribution - thái...

4 6 0 FAMILIES AND MAPS

class A c t i n o p t e r y g i i

Subclass N e o p t e r y g i i

Order Perciformes; Suborder Gobioidei

(per) Famdy O d o n t o b u t i d a e — o d o n t o b u t i d s [ o - d o n ' - t o - b L V - t i - d e ]

HOESE A N D GILL (1993) ERECTED THE FAMILY Odontobutidae for three genera that were formerly contained within the Eleotridae. Odontobutids are found in fresh waters of the northwest Pacific, including China, Korea, Japan, Russia, northern Vietnam, northeastern Thailand, and Laos (Iwata et al., 1985; Hoese and Gill, 1993; Sakai et al., 1996; Vidthayanon, 1995; Kottelat, 1998). Two specimens have been reported from the Philippines at Zamboanga (Herre, 1927) but have not yet been confirmed. Three plesiomorphic characters are shared with the Rhyacichthyidae and synapomorphies involving the pectoral girdle, dorsal f in , and scale morphology suggest monophyly of a group consisting of all gobioids except rhyacichthyids and odontobutids (Hoese and Gill , 1993). Although Hoese and Gill utilized a characteristic of the caudal skeleton—specifically the procurrent cartilages—as an important feature in establishing the Odontobutidae, it

ACTINOPTERYGIANS—RAY-FINNED FISHES

FIGURE 292. Odontobutis obscura (reproduced with permission from Weber and DeBeaufort, 1953, Fig. 75).

occurs only in Micropercops. The morphology of the procurrent cartilages in Odontobutis and Percottus is unremarkable (R. E. Watson, personal communication).

There are about eight species in the genera Micropercops, Odontobutis, and Perccottus (Hoese and Gill , 1993; Saki et al., 1996; Vidthayanon, 1995). The body form is that of an eleotrid. The lateral line is absent, and the pelvic fins are separate and do not form a sucking disk. The caudal fin is rounded. Nichols (1943) listed Micropercops cinctus, M. swinhonis, and M. dabryi from eastern China. Masuda et al. (1984) recorded Odontobutis obscura (Fig. 292) from central and southern Japan. Odontobutis obscura, with several subspecies, occurs widely throughout the range of the family (Iwata et al., 1985). Sakai et al. recorded O. interrupta and O. platycepbala from Korea. Odontobutis aurarmus was recently described from northeastern Thailand (Vidthayanon, 1995) and O. aspro from Laos (Kottelat, 1998). Berg (1948/1949) included Perccottus glehni from the Amur basin, China, and northeast Korea.

Map references: Berg (1948/1949), Herre (1927), Hoese and Gill (1993), Iwata et al. (1985),* Kottelat (1998), Sakai et al. (1996),* Vidthayanon (1995)*

Class A c t i n o p t e r y g i i



(per) Family E l e o t r i d a e — s l e e p e r s , g u d g e o n s [ e - l e - o ' - t r i -

ELEOTRIDS ARE FOUND IN TROPICAL, SUBTROPICAL, AND occasionally temperate, marine, brackish, and fresh waters throughout the world. They form an important component of the freshwater fauna of Australia, New Guinea, New Zealand, Hawaii, and islands of the

E l e o t r i d a e


Indo-Pacific. There are approximately 35 genera and 150 species of sleepers (Nelson. 1994). More of these species occur in fresh water than in brackish or marine waters.

The pelvic fins of eleotrids are separate and do not form a sucking disc as do the fins of most gobies. The mouth of sleepers is usually upturned or terminal but never inferior (Allen, 1991). The body may be covered with cycloid or ctenoid scales. There are no lateral line canals along the body.

The two eleotrid subfamilies recognized by Nelson (1994) (Butinae and Eleotrinae) are treated as subfamilies within the Gobiidae by Hoese and Gill (1993). The Butinae is a tropical group of 13 genera found in estuaries and freshwater streams of the Indo-Pacific and west Africa. The Eleotrinae is cosmopolitan in fresh water and mangrove habitats. It contains 22 genera.

Many sleepers are colorful and make excellent aquarium animals. They are easy to breed in captivity. Eleotrids attach their eggs to the substrate or vegetation. It is thought that the young of most freshwater eleotrids are swept downstream into brackish waters, and that the juveniles eventually migrate back into fresh waters to complete their life cycle. This suggests a marine origin for the family. Many eleotrids are benthic, but a large number have well-developed swim bladders. Al l eleotrids are carnivorous and feed primarily on insects, crustaceans, and small fishes. The common name "sleeper" reflects the fact that these fish spend a great deal of time resting on the bottom or positioned parallel to a stick or other object waiting for a potential meal. However, they can move very quickly and stop suddenly when chasing prey.

The fat sleeper, Dormitator maculatus, occurs in streams along the Atlantic and Gulf coasts of North America from as far north as the Hudson River of New York through the West Indes to Brazil (Lindquist, 1980a). This species approaches 60 cm TL and is used as a food fish. The largest sleeper, and probably the largest gobioid species, is Oxyeleotris marmorata at 66 cm TL (Roberts, 1989a). This species is from fresh water in Thailand, the Malay Peninsula, Sumatra, and Borneo and is considered a delicacy. Most eleotrids are much smaller at about 30-100 mm TL.

Gery (1969) wrote that eleotrids enter, more or less occasionally or for spawning, into South American rivers. Tropical America has four genera restricted to fresh waters: Microphilypnus (Amazon), Leptophilypnus (Atlantic and Pacific slopes of Central America), Hemieleotris (Pacific slope from Costa Rica to Colombia), and Gobiomorus (rivers of the Antilles and Central and South America) (Banarescu, 1990).

Allen (1991) recorded 41 species of gudgeons (as eleotrids are called in Australia and New Guinea) from the fresh waters of New Guinea. Morgunda is a prominent part of the freshwater fish fauna of New Guinea and Australia, with 16 species recorded from New Guinea and at least 6 species from Australia. Half of the New Guinean species are from the Lake Kutubu region in the Southern Highlands of Papua New Guinea.

4 6 4

FAMILIES AND MAPS

FIGURE 293. Hypseleotris galii as Carassiops gain (Waite, 1904a, Plate XXXTV, Fig. 2).

Oxyeleotris fimbriata is widely distributed throughout New Guinea on both sides of the central divide (Allen, 1991).

In Australia, Hypseleotris is an important component of the freshwater fish fauna, with about 9 of the 25 eleotrid species (Allen, 1989). Hypseleotris galii (Fig. 293) occurs in coastal streams from Fraser Island, Queensland, to Eden in southern New South Wales (Larson and Hoese, 1996). The species is very small, but males (55 mm SL) are larger than females (40 mm) and develop a hump on the head behind the eyes during breeding season. Normally gray or bronze, breeding males may become black with orange fins. Hypseleotris klunzingeri is widespread in streams and ponds of the Murray-Darling system of eastern Australia as well as in coastal drainages from the Fitzroy River in central Queensland to the Hunter River of New South Wales (Larson and Hoese, 1996). It often forms dense schools in midwater. It prefers vegetated habitats. Maximum TL is only about 45 mm. Hypseleotris compressa from coastal streams around the northern two-thirds of Australia and south-central New Guinea is a beautiful, little, sexually dimorphic fish. The male's dorsal and anal fins become intensely blue, black, and orange during the breeding season, and the ventral surface also develops a bright orange coloration. They readily breed in captivity, and the males guard the nest. The female may deposit up to 3000 eggs (Merrick and Schmida, 1984). Maximum TL is 100 mm, but most specimens are only about half this size.

Philypnodon grandiceps (Fig. 294) occurs throughout the Murray-Darling River system and coastal stream of southeastern Australia, including the north coast of Tasmania. It can reach 115 mm TL. Mogumda adspera lives in Pacific coast drainages from Cape York peninsula southward to the Clarence River of northern New South Wales. Mogumda mogumda occurs in freshwater streams of northern Australia from the Kimberley region to Cape York peninsula and in central southern New Guinea. Mogumda larapintae is found in the Finke River system of central Australia. Other desert Mogumda remain to be described (G. Allen, personal communication). Milyeringa Veritas is an eyeless, white or pinkish


FIGURE 294. Philypnodon grandiceps (Waite, 1904a, Plate XXXVI, Fig. 2).

gudgeon from wells and sink holes of the North West Cape region of Western Australia. Two species of Kimberleyeleotris are known only from the Drysdale and Mitchelle Rivers of Western Australia. Other genera found in Australia include Butis and Gobiomorphus.

Eleotrids are called bullies in New Zealand, and they compose a significant part of a depauperate freshwater fish fauna. There are six species of Gobiomorphus in New Zealand's rivers, and all occur on both North and South Island (McDowall, 1990). Gobiomorphus basalis is only known from one population in the alpine reaches of the Awatere River system on South Island, but it is relatively widespread throughout the North Island. Gobiomorphus huttoni (Fig. 295) is the only eleotrid to occur on Stewart Island at the southern tip of South Island. This is the most southerly occurrence of any eleotrid. This colorful species is common throughout New Zealand. Gobiomorphus cotidianus is the most widespread bully in New Zealand. Gobiomorphus gobioides is the largest New Zealand bully at 22 cm TL.

In Hawaii, the endemic Eleotris sandwicensis is one of the few prominent stream fishes (Gosline and Brock, 1965). Kottelat et al. (1993) list 10

FIGURE 295. Gobiomorphus huttoni, female (reproduced with permission from McDowall, 1990, Fig. 18.1 A).

466

FAMILIES AND MAPS

genera and 19 species from the islands of Indonesia including Sulawesi. Herre (1927) presented an account of 18 genera and 32 species of eleotrids from the Philippines. Talwar and Jhingran (1992) reported Eleotris, Butis, Ophiocara, Ophieleotris, and Odonteleotris from fresh waters in the Indian region. Ophiocara porocephala (Fig. 296) reaches 3 cm TL and enters fresh waters throughout the Indo-Pacific region (Talwar and Jhingran, 1992).

FIGURE 296. Ophiocara porocephala as Eleotris porocephalus (Day, 1878b, Plate LXVTJ, Fig. 1).

FIGURE 297. Kribia kribensis as Eleotris kribensis (Boulenger, 1916, Fig. 9).

There are 11 genera and 25 species of Eleotridae recorded from Africa (Mauge, 1986c). Kribia is an African genus restricted to fresh water. Kribia kribensis (Fig. 297) is found from Guinea to the Congo, K. leonensis is restricted to Sierra Leone, K. nana occurs in Lake Chad and the Congo River system, and K. uelensis is endemic to the Uelle River basin of central Africa (Harrison and Miller, 1992a; Mauge, 1986c). These species are less than 50 mm TL. Eleotris, Dormitator, and Hypseleotris also occur in African fresh waters. Ratsirakia and two species of the subterranean Typhleotris are restricted to fresh waters of Madagascar (Stiassny and Raminosoa, 1994).

Map references: Allen (1989, 1991),* Bartholomew (1911),* Berg (1948/1949), Bussing (1998),* Eschmeyer and Herald (1983), Harrison and Miller (1992a),* Lindquist (1980),* McDowall (1990),* Masuda et al. (1984), Mauge (1986c), Pethiyagoda (1991),* Skelton (1993),* Talwar and Jhingran (1992)


Class A c t i n o p t e r y g i i



(per) Family G o b i i d a e — g o b i e s ( g o - b i ' - i - d e ) [ g o - b e ' - i - d e ]

THE GOBIIDAE, AS CONSTITUTED ACCORDING TO NELSON (1994), is the largest marine fish family with about 212 genera and approximately 1875 species. If one were to include the eleotrids, as recommended by Hoese and Gill (1993), the number of species would exceed 2000 and the Gobiidae would rival the Cyprinidae as the largest fish family (or vertebrate family) in the world. The Cichlidae would not be far behind. The cyprinids are fairly well-known, but there are many more marine and estuarine gobies yet to be described. The Gobiidae is the marine family with the largest number of recently described new species (Berra, 1997a; Berra and Berra, 1977).

Gobies are distributed throughout the world in marine, brackish, and fresh water, primarily in the tropics and subtropics and especially in the Indo-West Pacific, but marine species are known from the subarctic of Norway and Iceland and subarctic streams of southern Siberia. Gobiids, like eleotrids, are common in freshwater streams of islands. Gobies usually have united pelvic fins that form a sucking disc, and all freshwater species have united pelvic fins. Some reef gobies may have separate pelvics. However, the degree of separation of gobiid pelvic fins is highly variable, and it is not always easy to distinguish an eleotrid from a gobiid. There are two dorsal fins. They lack a lateral line on the body, and scales may be cycloid, ctenoid, or absent. Most gobies are small under 10 cm, and their small size makes identification difficult. The smallest known vertebrate is a marine goby, Trimmatom nanus, from the Chagos archipelago in the Indian Ocean. This minute, scaleless fish is sexually mature at about 8-10 mm SL (Winterbottom and Emery, 1981; Kottelat and Vidthayanon, 1993), but an even smaller genus is being described from southern Japan (R. E. Watson, personal communication). In the Philippines, two freshwater species, Pandaka pygmaea and Mistichtbys luzonensis, are nearly as tiny at 10 or 11 mm. Some species, such as Gobioides broussenetii from the Caribbean, may reach 50 cm TL.

With such a large number of species comes a great diversity of lifestyles, especially in marine gobies. Some gobies (Gobiosoma) function as cleaner fishes that remove external parasites from coral reef fishes (Limbaugh, 1961; Losey, 1987). Other gobies (Crytocentrus steinitzni) have a symbiotic relationship with shrimp and share a burrow with them (Preston, 1978;

G o b i i d a e

469


Karplus, 1979). Other gobies live in contact with sponges, corals, and sea urchins.

Mudskippers (Boleophtbalmus, Periophthalmus, Periophthalmodon, and Scartelaos) are amphibious, euryhaline gobies that leave the water and dart about on mudflats in search of food such as insects and crustaceans. Some respiration takes place cutaneously and via water retained in the branchial chambers. Their eyes are on moveable short stalks and are adapted for seeing in air as well as in water. They can even climb the prop roots of mangrove trees. See Polunin (1972) and Nursall (1981) for photographs and descriptions of their remarkable behavior. Murdy (1989) provided a taxonomic revision of mudskippers and their relatives (Oxudercinae). Consult Graham (1997) for an extensive review of mudskipper adaptations for aerial respiration. Horn et al. (1999) reviewed the biology of intertidal fishes.

Hoese (1984) subdivided the Gobiidae into four subfamilies, and this arrangement was followed by Birdsong et al. (1988). Pezold (1993) added a f i f th subfamily. These are Oxudercinae, Amblyopinae, Sicydiinae, Gobionellinae, and Gobiinae. The Sicydiinae comprise freshwater gobies (larvae are euryhaline), and the Gobionellinae includes mostly estuarine gobies as well as many species inhabiting fresh waters. The total number of goby species in fresh water is not known, but it is approximately 200 (Hoese, 1994). Of these, about 100 species are sicydi-ine gobies that make up a group of tropical and subtropical stream fishes that exhibit a high degree of island endemism, especially in the Pacific (Parenti and Maciolek, 1993, 1996). These amphidromous gobies spend most of their life in fresh water, where they breed. The larvae are swept downstream to the sea, where they grow before ascending streams. Most Pacific islands with sufficient elevation to maintain perennial stream systems are populated by sicydiine gobies, as are high-gradient, coastal streams.

Currently, the six genera of the Sicydiinae are Cotylopus, Lentipes, Sicydium, Sicyopterus, Sicyopus, and Stiphodon (Watson, 1995; Parenti and Maciolek, 1996). New genera from Vanuatu and west Africa remain to be described (R. E. Watson, personal communication). Cotylopus is only known from mountainous streams of Reunion and is retained as a synonym of Sicyopterus in most accounts (Watson, 1995). Sicydium occurs in insular and coastal streams of the tropical eastern Pacific and the Atlantic, including the Caribbean Sea. The other genera occur in the central Pacific. According to Parenti and Maciolek (1996), Sicyopus is the most plesiomor-phic genus, and Sicyopterus is one of the most derived sicydiines. Both genera also inhabit the Indian Ocean. The polarization of genera as stated by Parenti and Maciolek (1993) is challenged by Watson based on teeth characteristics of Sicyopus (Watson, 1999a, personal communication). Parenti and Thomas (1998), using pharyngeal jaw morphology, explored hypotheses of relationships of sicydiine gobies. Stiphodon from various

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