Gobiesocoidei

(Clingfishes and singleslits)

Class Actinopterygii

Order Perciformes

Suborder Gobiesocoidei

Number of families 1

Evolution and systematics

No clingfishes and singleslits have been identified from the fossil record, and the evolutionary history remains to be examined with molecular methods. The examination of morphological relationships has led to the formulation of competing hypotheses to describe the history and systematic position of clingfishes and singleslits, and these questions are far from resolved. Some workers place these fishes in the order Gobiesociformes along with the dragonets (Callionymidae) and draconetts (Draconettidae). Others place the Gobiesocidae in the suborder Gobiesocoidei within the Perciformes, next to the suborder Callionymoidei (Callionymidae and Draconettidae); such is the treatment in this chapter.

The family Gobiesocidae is partitioned into two subfamilies, the Gobiesocinae (clingfishes) and the Cheilobranchinae (singleslits) within the Gobiesocoidei. Both subfamilies are united by the presence of a joint that connects the supraclei-thrum and cleithrum bones within the pelvic girdle. Two other specializations link these two subfamilies: a joint connecting the interopercle and epihyal and a heart with a structure unique to both subfamilies. The singleslits are sometimes recognized as a separate family within the suborder Gobiesocoidei; the family consists of a single genus, Alabes, with four species. Briggs recognized the following subfamilies (denoted here as tribes), which now are placed under the Gobiesocinae: the primitive Trachelochismini, the monotypic Haplocylicini, the Lepadogastrini, the monotypic Chorisochismini, the Diplocrepini, the Gobiesocini, and the highly specialized Diademichthyini, which are mostly commensal with sea urchins and crinoids. Briggs concluded that more specialized taxa are found in the tropical Indo-West Pacific region; intermediate taxa are found in the eastern Pacific, western Atlantic, and eastern Atlantic; and relict Southern Hemisphere taxa are confined to the cool temperate waters of southern Africa, Australia, and New Zealand. There are at least 120 species of clingfishes (two with two subspecies each and one with three subspecies) in 44 genera plus undescribed genera and species.

The known genera include Acyrtops (two species), Acyrtus (two species), Alabes (four species), Apletodon (four species, one with two subspecies), Arcos (five species), Aspasma (one species), Aspasmichthys (one species), Aspasmodes (one species), Aspasmogaster (four species), Chorisochismus (one species), Cochleoceps (four species), Conidens (two species), Creocele (one species), Dellichthys (one species), Derilissus (two species), Diademichthys (one species), Diplecogaster (three species, one with three subspecies), Discotrema (one species), Eckloniaichthys (one species), Gastrocyathus (one species), Gastrocymba (one species), Gastroscyphus (one species), Gobiesox (26 species), Gouania (one species), Haplocylix (one species), Kopua (one species), Lecanogaster (one species), Lepadichthys (nine species), Lepadogaster (three species, one with two sub-species), Liobranchia (one species), Lissonanchus (one species), Modicus (two species), Opeatogenys (two species), Parvicrepis (one species), Pherallodichthys (one species), Pherallodiscus (one species), Pherallodus (two species), Posidonichthys (one species), Propherallodus (one species), Rimicola (five species), Sicyases (two species), Tomicodon (13 species), and Trachelochismus (two species).

Physical characteristics

The clingfishes are denoted mainly by the presence of pelvic fins modified into a sucking disk. Unlike the gobies (Gobiidae, Perciformes), which possess pelvic fins modified into a suction cup, the clingfishes lack a spinous dorsal fin. With some exceptions, clingfishes are tadpole-like, with broad, depressed heads and flattened bodies. Some species, however, are elongate with pointed snouts. Clingfishes lack scales but shield themselves with a mucus that forms a protective coat around their bodies. The lateral line is well developed anteriorly but is either small or missing posteriorly.

There is a well-developed network of sensory pores on the head. There is one dorsal and one anal fin.

The sucking disc is of two types: a single disc, in which the anterior and posterior components form a single continuous structure, and a double disc, in which the larger anterior and smaller posterior components form two discs separated by a single wall or edge. The pectoral fins generally are small and are positioned posterior to the sucking disk. The dorsal nostril and, usually, the posterior nostril are both tubular. The swim bladder is absent. Color patterns are variable and range from black to orange, brown, green, or red. Additionally, there may also be contrasting stripes, bars, or spots of yellow, blue, green, brown, gray, or white. The larvae are elongate, torpedo-shaped, and laterally compressed, and they become more rounded with growth. The gas bladder, located above the gut, is present but is lost after settlement. Body sizes typically are small, less than 1.9–2.4 in (5–6 cm) in total length, but at least one temperate species from southern Africa, the rocksucker (Chorisochismus dentex), reaches 11.8 in (30 cm). Clingfishes are sexually dimorphic, with differences in the size of the urogenital papillae and also in body size—males generally are larger than females. Two species reportedly produce a skin toxin.

The singleslits are eel-like in shape, with diminutive pelvic fins and a vestigial sucking disc. Their dorsal and anal fins are modified to resemble those of true eels. Soft rays are present only in the caudal fin. The posterior nostril is absent in one species. Color patterns are also variable and range from translucent pink or green to light green, brown, gray, or black. Spots or bars of black, gray, brown, or yellow may also be present. Usually they are less than 1.9 in (5 cm) in length, but one species, the common shore-eel (Alabes dorsalis) of Australia, reaches 4.7 in (12 cm).

Distribution

Clingfishes occur in marine and brackish waters in tropical, warm temperate, and temperate zones of the Atlantic, Indian and Pacific Oceans. General distribution patterns may be described best by relying upon Briggs' classification scheme, as modified here. Thus, the Trachelochismini consists of Southern Hemisphere relict species restricted, except for two Northern Hemisphere species, to temperate Australia and New Zealand. The monotypic Haplocylicini is endemic to New Zealand. Members of the Lepadogastrini occur in the eastern Atlantic and Mediterranean Sea, with one species found in deeper waters of southern Africa. The Chorisochismini is monotypic and also is found in southern Africa, but in shallower, warm temperate waters. The Diplocrepini is distributed across the Indo-West Pacific, with one deepwater relict species occurring in the Caribbean. The Gobiesocini occurs in the western Atlantic and eastern Pacific; some also are found in freshwater streams of the Caribbean, Central America, and Cocos Island in the eastern Pacific. One monotypic genus occurs in southern Africa. The Diademichthyini also is found in tropical waters of the Indo-West Pacific region. Singleslits are restricted to temperate inshore waters of the southern Australian mainland, Tasmania, and Norfolk Island.

Habitat

Clingfishes are largely inshore fishes adapted to clinging to the substrate. Thus, they are able to colonize high-energy habitats and withstand breaking waves and surging waters. Many species occur on or under boulders and rocks, in crevices, and on rocky slopes and rock faces. Species adapted to living in tide pools are capable of remaining out of water for a number of days, if they are kept moist and out of sunlight. Other species are found in close association with corals, soft corals, sponges, and ascidians. Some species live among crinoids or sea urchins. The crinoid clingfish, Discotrema crinophila, lives secluded among the arms of crinoids. Diademichthys lineatus has strayed from the general clingfish body plan by evolving an elongated shape that nearly mimics the long spines of Diadema sea urchins. Other species cling to the blades of sea grasses and to seaweed and kelp. Some members of the genus Gobiesox occur in freshwater streams. Singleslits also live in inshore marine waters, usually in shallow tide pools and often under stones or in close association with seaweed, the stems or branches of which they may cling to or wrap around with their prehensile caudal fins and flexible bodies.

Behavior

Clingfishes and singleslits tend to be secretive, although the reef-associated species commensal with sea urchins and other organisms are more mobile and swim in the water column. Territoriality has been reported for numerous species and is suspected for the rest. This trait is more pronounced in males. Most observations of other behavioral activity have been confined to aquarium studies; field investigations, where physically possible, are desirable.

Feeding ecology and diet

The clingfishes generally feed on smaller benthic invertebrates, but there are significant exceptions to this rule. Cling-fish diets, depending upon species, body size, and habitat, include various algae, crustaceans (including amphipods, copepods, small crabs and shrimps), polychaete worms, small bivalves, limpets and other gastropods, chitons, body parts of host sea urchins, and small fishes. Members of the genus Cochleoceps are cleaner fishes that remove ectoparasites from boxfishes, porcupinefishes, morwongs, and other species. The singleslits feed mainly on small benthic invertebrates. Cling-fishes are preyed upon by other fishes, shorebirds, and possibly crabs living in the intertidal zone.

Reproductive biology

Reproductive behavior and ecology have been described for a number of species, mainly from aquarium studies, and, as expected, there is species-specific variation around a common theme. Courtship is paired and initiated by the male within his territory. Generally, the male nudges the female's abodomen; if the female responds positively, he moves parallel to her flank and undulates or quivers. After some time, the female will undulate her body and lay demersal eggs singly until a small clutch is produced. (Demersal eggs are those that are deposited upon the substrate, such as the sea floor.) Eggs may be laid on stones, algae, or other substrates. Depending upon the species, egg laying may last from several minutes to a few hours. The male fertilizes the eggs during this process, and, upon completion, the clutch either is guarded by the male or is abandoned by the pair. There is some evidence of mating by internal fertilization, with female parental care, in the weedsucker (Eckloniaichthys scylliorhiniceps). This species is a monotypic clingfish endemic to South Africa that may share this trait with a few other species. Spawning or mating may be seasonal, during warmer months, at higher latitudes. At tropical or subtropical latitudes, spawning or mating may take place year-round or seasonally if water temperatures become either too warm or too cold. Larvae of all species are probably planktonic.

Conservation status

No species is currently on the IUCN Red List. Marine and freshwater species with limited distributions may be at risk from habitat destruction or pollution.

Significance to humans

Some species are collected for the aquarium trade.

Species accounts

List of Species

Common shore-eel

Alabes dorsalis

family

Gobiesocidae

taxonomy

Alabes dorsalis Richardson, 1845, attributed to the northwest coast of Australia, but this is likely a type locality error.

other common names

English: Shore eel.

physical characteristics

Body small and elongate (eel-like) with reduced fins. Vestigial sucking disc. Color variously gray to black, with prominent green spots along the flank and gray to black blotches regularly spaced along the dorsal fin. Grows to 4.7 in (12 cm) in length.

distribution

Australia from southern New South Wales south to Tasmania and west to central South Australia.

habitat

Intertidal zone under rocks.

behavior

Poorly known. This species is secretive and seldom seen. The common shore-eel moves rather like a true eel among and under rocks, into holes, and along the bottom of tide pools. Nothing is known about its social behavior, although it may be

territorial, like others in its family. May tolerate exposure to air during exceptionally low tides.

feeding ecology and diet

Feeds upon small benthic invertebrates.

reproductive biology

Unknown. Spawning probably is seasonal, during warmer months, and paired with the deposition of demersal eggs. Larvae are probably planktonic.

conservation status

Not listed by the IUCN. May be at risk from habitat destruction or pollution.

significance to humans

May be collected infrequently for aquaria.

Crinoid clingfish

Discotrema crinophila

family

Gobiesocidae

taxonomy

Discotrema crinophila Briggs, 1976, between Stuart and Yanuca islands, Benga Lagoon, Benga, Fiji Islands.

other common names

None known.

physical characteristics

Elongate body, with a rounded snout. There are 8–9 dorsal fin soft rays and 7–8 anal fin soft rays. Color is black, with a yellow stripe that runs along each flank from the snout to the base of the caudal fin and a mid-dorsal yellow stripe that joins the lateral stripes at the snout.

distribution

Western Pacific, from the Ryukyu Islands of southern Japan south to Taiwan, southeast to the Bismarck Archipelago in Papua New Guinea, and east to Fiji and Christmas Island, Line Islands (Kiribati). Specimens reported from the Great Barrier Reef, and possibly the Bismarck Archipelago, represent an undescribed species.

habitat

Lives among the arms of crinoids on coral reefs at a depth range of 26–66 ft (8–20 m).

behavior

Poorly known. Swims about the arms of crinoids.

feeding ecology and diet

Feeds upon small invertebrates.

reproductive biology

Poorly known but likely lays demersal eggs, perhaps at the base of a crinoid.

conservation status

Not listed by the IUCN.

significance to humans

May be collected infrequently for aquaria.

Eastern cleaner-clingfish

Cochleoceps orientalis

family

Gobiesocidae

taxonomy

Cochleoceps orientalis Hutchins, 1991, Big Island, off Wollon-gong, New South Wales, Australia.

other common names

None known.

physical characteristics

Small, tadpole-shaped body with rounded caudal fins and a moderately sized sucker disc. There are 5–6 dorsal fin soft rays and 4–6 anal fin soft rays. Color pattern is tiny brown to red spots on an orange to yellow or greenish-yellow background; irridescent blue dashes, lines, or spots occur on the dorsum. Grows to 2.2 in (5.5 cm) in length.

distribution

Southwest Pacific from central New South Wales south to eastern Victoria, Australia.

habitat

Occurs on sponges or ascidians on deeper temperate rocky reefs and on kelp or seaweed in shallower subtidal habitats. Found as deep as 33 ft (10 m).

behavior

In addition to its cleaning behavior, this species probably is territorial.

feeding ecology and diet

A cleanerfish, it feeds upon ectoparasites plucked from various species of fishes.

reproductive biology

Poorly known, but females deposit eggs on algae within a male's territory, and both parents provide care.

conservation status

Not listed by the IUCN.

significance to humans

May be collected infrequently for aquaria.

Northern clingfish

Gobiesox maeandricus

family

Gobiesocidae

taxonomy

Gobiesox maeandricus Girard, 1858, San Luis Obispo, California, United States.

other common names

None known.

physical characteristics

There are 13–16 dorsal soft rays; 12–14 anal soft fin rays; a rounded caudal fin; and short, rather broad pectoral fins with a fleshy palp that bridges between the base of the fin and the gill opening. The color pattern consists of a light olive-brown or cherry-red background with either dark reticulations or mottled shades of lighter color and a series of white bars between the eyes. In juveniles the white bars also run along the back and on the edge of the caudal fin. Grows to about 6.3 in (16 cm) in length. Males tend to be larger than females.

distribution

Eastern Pacific from southeastern Alaska south to southern California; also found at Revillagigedo Island off Baja California, Mexico.

habitat

Lives among algae or under rocks in the intertidal zone and in the upper canopy of kelp forests, to a depth of 26 ft (8 m). The ability to breathe atmospheric air with its gills allows it to tolerate exposure, if it is kept moist under rocks or algae.

behavior

Males maintain territories. Both males and females tend to be somewhat secretive, clinging to surfaces and moving only to feed or interact with others.

feeding ecology and diet

Feeds upon small mollusks and crustaceans found on rocks, algae, and kelp.

reproductive biology

As with others in this genus, this species likely spawns demersal eggs in a nest prepared by a male within his territory.

Spawning may take several hours. Males provide parental care. The larvae are planktonic.

conservation status

Not listed by the IUCN.

significance to humans

Collected mainly for public aquaria.

Pygmy shore-eel

Alabes parvulus

family

Gobiesocidae

taxonomy

Alabes parvulus McCulloch, 1909, rock pools near Sydney, New South Wales, Australia.

other common names

None known.

physical characteristics

Elongate and eel-like body with reduced fins. Vestigial sucking disc. Grows to 1.9 in (5 cm) in length.

distribution

Temperate waters of southern Australia and Norfolk Island.

habitat

Inhabits the intertidal zone. Found in tide pools, in brown seaweed beds and under rocks, to a depth of 20 ft (6 m).

behavior

Poorly known. Has been found in small groups of three to six individuals, and social interactions may be expected to take place regularly.

feeding ecology and diet

Feeds upon small benthic invertebrates.

reproductive biology

Largely unknown. Spawning is probably seasonal, during warmer months, and is paired with the deposition of demersal eggs. Larvae are probably planktonic.

conservation status

Not listed by the IUCN.

significance to humans

May be collected infrequently for aquaria.

Rocksucker

Chorisochismus dentex

family

Gobiesocidae

taxonomy

Chorisochismus dentex Pallas, 1769, type locality not specified.

other common names

None known.

physical characteristics

Large, tadpole-like body with a very broad head. There are 7–9 dorsal fin soft rays, 6–7 anal fin soft rays, 21–24 pectoral fin soft rays, and 8–10 caudal fin rays. The teeth are large and conical in shape. Perhaps the largest of all clingfishes, this species reaches 11.8 in (30 cm) in length.

distribution

Southeast Atlantic, from Namibia south to northern Natal in South Africa.

habitat

Tide pools and rocks and boulders in the intertidal zone.

behavior

Poorly known. Males likely are territorial. Owing to its large size, rocksuckers are probably more mobile than other demersal clingfishes.

feeding ecology and diet

A carnivore that preys upon limpets and sea urchins. This species uses its large upper incisiform teeth to pry limpets off rocks. The undigested shell fragments are passed through the gut and emerge in mucous capsules.

reproductive biology

Poorly known. Females probably lay demersal eggs in a male's territory, and the male performs parental care. The larvae are probably planktonic.

conservation status

Not listed by the IUCN.

significance to humans

May be collected infrequently for aquaria.

Sonora clingfish

Tomicodon humeralis

family

Gobiesocidae

taxonomy

Tomicodon humeralis Gilbert, 1890, Puerto Refugio (Angel Island) and La Paz, Mexico.

other common names

Spanish: Chupapiedra de Sonora.

physical characteristics

Compressed, elongate body with a broad head and a large sucking disc. There are 8–9 dorsal fin soft rays, 6–7 anal soft fin rays, 17–19 pectoral fin soft rays, and 9–10 caudal fin soft rays. The color pattern consists of light diagonal stripes along the entire body, with a pair of dorsal spots positioned just behind the head, about even with the pectoral fins. Grows to 3.3 in (8.5 cm) in length. Sexually dimorphic, with males larger than females.

distribution

Endemic to the Gulf of California from Punta Borrascoso to Guaymas in Sonora and from San Felipe to Cabo San Lucas in Baja California.

habitat

A common clingfish found mainly in the upper and mid-inter-tidal zones, usually under rocks, to which they cling, and in little or no water. If kept moist, this fish is able to withstand extreme temperatures when exposed to air.

behavior

This species is secretive, clinging to the undersides of rocks and moving over rocky surfaces to feed. Their movements are related to tidal movements. Activity is greater at high tide, and there is little or no activity at low tide. Males are territorial.

Both males and females secrete large amounts of mucus, which coats their bodies and protects them from desiccation.

feeding ecology and diet

A diurnal predator that feeds upon small crustaceans, including barnacles and limpids.

reproductive biology

Spawning begins in late spring or early summer and is paired with demersal courtship. A single male may mate with more than one female. Eggs are adhesive and laid on the underside of rocks, where they are guarded by the male, sometimes with the aid of one or more females. During low tide, the parents secrete mucus that protects the eggs from exposure. The larvae are planktonic.

conservation status

Not listed by the IUCN.

significance to humans

May be collected infrequently for aquaria.

Two-spotted clingfish

Diplecogaster bimaculata bimaculata

family

Gobiesocidae

taxonomy

Diplecogaster bimaculata bimaculata Bonnaterre, 1788, seas of England, although no types were designated.

other common names

English: Two-spotted sucker.

physical characteristics

Flattened and elongated body, with a somewhat triangular snout. The dorsal fin has 4–7 soft rays, and the anal fin has 4–7 soft rays. The gill opening is minute. Color varies but usually is red with yellow on the ventral surface and a pattern of blue and brown spots over the body surface. Males are distinguished by a purple or red spot, ringed in yellow, on each flank, behind the pectoral fins. Grows to 2.4 in (6 cm) in length.

distribution

Eastern Atlantic Ocean, from Norway and the Faroe Islands south to Gibraltar and the western Mediterranean and Adriatic. The subspecies Diplecogaster bimaculata euxinica is found in the Black Sea and D. bimaculata pectoralis from the offshore Canary and Cape Verde Islands and the Azores.

habitat

A temperate species found on sea grass beds, rocky bottoms, bivalve banks, and mud bottoms. Reported to favor shells. Depths range between 59 and 180 ft (18–55 m) in cooler waters and down to 328 ft (100 m) in the warmer Mediterranean.

behavior

Not well known. Males likely are territorial.

feeding ecology and diet

Feeds on small benthic invertebrates.

reproductive biology

Demersal courtship and spawning occur in the spring and summer months, with golden-colored eggs laid in masses under shells or stones. Males provide most of the parental care. Larvae are planktonic.

conservation status

Not listed by the IUCN.

significance to humans

Occasionally recovered from trawls working soft mud bottoms or bivalve banks.

Urchin clingfish

Diademichthys lineatus

family

Gobiesocidae

taxonomy

Diademichthys lineatus Sauvage, 1883, New Caledonia.

other common names

None known.

physical characteristics

Elongate and slender body, with spatulate snout and small fins. There are 13–15 dorsal fin soft rays, 12–14 anal fin soft rays, and 25–26 pectoral fin soft rays. Color is reddish or reddish brown, with paired yellow stripes and a yellow blotch on the caudal peduncle that extends to the caudal fin. Grows to 1.9 in (5 cm) in length.

distribution

From Oman and Mauritius in the western Indian Ocean east to Indonesia and Fiji, north to southern Japan, and south to northern Australia.

habitat

Inhabits shallow coral reefs among the long-spined sea urchins and branching corals, usually in holes or protected areas.

behavior

This clingfish swims, often in a dancing or undulating motion, between the spines of sea urchins, within coral heads, and in holes or small caves where sea urchins might be found. Their behavior is not known in any great detail and remains to be studied.

feeding ecology and diet

Takes burrowing bivalves on corals, eggs of shrimp commensal with sea urchins, and the tube feet of sea urchin hosts. Juveniles feed on the pedicellariae and sphaeridia of host urchins and on copepods and the eggs of shrimp that are commensal with sea urchins.

reproductive biology

This species probably courts and spawns in a manner similar to that of others in its family, except that courtship bouts most likely take place in the water column. The eggs are small and demersal, and the larvae are pelagic.

conservation status

Not listed by the IUCN. May be at risk from the loss of sea urchins because of overfishing, disease, or other factors and from habitat destruction and the effects of pollution on reefs.

significance to humans

May be collected for the aquarium trade.

Resources

Books

Bohlke, James E., and Charles C. G. Chaplin. Fishes of the Bahamas and Adjacent Tropical Waters. 2nd edition. Austin: University of Texas Press, 1993.

Briggs, J. C. "Clingfishes." In Encyclopedia of Fishes, edited by J. R. Paxton and W. N. Eschmeyer. San Diego: Academic Press, 1995.

Briggs, J. C., and J. B. Hutchins. "Clingfishes and Their Allies." In Encyclopedia of Fishes, 2nd edition, edited by J. R. Paxton and W. N. Eschmeyer. San Diego: Academic Press, 1998.

Eschmeyer, W. N., ed. Catalog of Fishes. 3 vols. San Francisco: California Academy of Sciences, 1998.

Helfman, G. S., B. B. Collette, and D. E. Facey. The Diversity of Fishes. Malden, MA: Blackwell Science, 1997.

Kuiter, R. H. Guide to Sea Fishes of Australia. Sydney: New Holland, 1997

Leis, Jeffrey M., and Brooke M. Carson-Ewart, eds. The Larvae of Indo-Pacific Coastal Fishes. Boston: Brill, 2000.

Lythgoe, J., and G. Lythgoe. Fishes of the Sea: The North Atlantic and Mediterranean. London: Blandford Press, 1991.

Masuda, H., K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino, eds. The Fishes of the Japanese Archipelago. Tokyo: Tokai University Press, 1984.

Myers, R. F. Micronesian Reef Fishes: A Field Guide for Divers and Aquarists. 3rd edition. Barrigada, Guam: Coral Graphics, 1999.

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

Randall, J. E., G. R. Allen, and R. C. Steene. Fishes of the Great Barrier Reef and Coral Sea. Honolulu: University of Hawaii Press, 1996.

Smith, M. M., and P.C. Heemstra, eds. Smiths' Sea Fishes. Berlin: Springer-Verlag, 1986.

Thomson, Donald A., Lloyd T. Findley, and Alex N. Kerstitch. Reef Fishes of the Sea of Cortez. 2nd edition. Tucson: University of Arizona Press, 1987.

Thresher, R. E. Reproduction in Reef Fishes. Neptune City, NJ: T.F.H. Publications, 1984.

Periodicals

Briggs, J. C. "New Species of Rimicola from California." Copeia 2002 (2002): 441–444.

Gosline, W. A. "A Reinterpretation of the Teleostean Fish Order Gobiosociformes." Proceedings of the California Academy of Sciences series 4, 37 (1970): 363–382.

Organizations

IUCN/SSC Coral Reef Fishes Specialist Group. International Marinelife Alliance-University of Guam Marine Laboratory, UOG Station, Mangilao, Guam 96913 USA. Phone: (671) 735-2187. Fax: (671) 734-6767. E-mail: [email protected] Web site: <http://www.iucn.org/themes/ssc/sgs/sgs.htm>

Terry J. Donaldson, PhD