Percoidei IV

(Goatfishes, butterflyfishes, angelfishes, chubs, and relatives)

Class Actinopterygii

Order Perciformes

Suborder Percoidei

Number of families 18

Evolution and systematics

The suborder Percoidei contains more than 70 families and 2,800 species. This section on the Percoidei deals with 18 families of fishes. The families are the Mullidae (goatfishes), Toxotidae (archerfishes), Dichistiidae (galjoens), Kyphosidae (sea chubs or rudderfishes), Paracorpididae (jutjaws), Drepaneidae (sicklefishes), Monodactylidae (monos or moonfishes), Chaetodontidae (butterflyfishes), Pomacanthidae (angelfishes), Enoplosidae (oldwives), Pentacerotidae (armorheads and boarfishes), Nandidae (leaffishes), Oplegnathidae (knifejaws), Cirrhitidae (hawkfishes), Chironemidae (kelpfishes), Aplodactylidae (seacarps or marblefishes), Cheilodactylidae (morwongs), and Latridae (trumpeters).

Fossil records indicate that perciform fishes date back as far as the Cenozoic era (about 65 million years ago), with percoids evolving rapidly during the Eocene epoch. This rapid evolution is evident in such families as the Chaetodontidae, Pomacanthidae, Mullidae, Kyphosidae, and Cirrhitidae, which contain the most species in the group. A number of families are placed in groups with variable degrees of formalization. For example, the Squammipinnes is an unranked group, denoted by having rows of scales covering the base of the dorsal and anal fins, and includes the Monodactylidae, Toxotidae, Chaetodontidae, and Pomacanthidae. The Kyphosidae consists of four subfamilies, three of which were recognized previously as separate families. (A new molecular study of this polyphyletic group suggests that the subfamilies, which appear to be closely related to the Kuhlidae, might be recognized again as separate families.) The Nandidae also consists of three subfamilies recognized on the basis of their geographical distribution, which parallels the breakup of Gondwana (the supercontinent, made up of South America, Africa, Antarctica, India, and Australia), thus making the nandids an ancient group. Leaffishes have been linked falsely with labyrinth fishes of the suborders Anabantoidei (gouramies and allies) and Channoidei (snakeheads). The superfamily Cirrhitoidea is denoted by the presence of pectoral fins with five-to-eight unbranched and elongated lower fin rays on the pectoral fin. Members include the Cirrhitidae, Chironemidae, Aplodactylidae, Cheilodactylidae, and the Latridae.

Physical characteristics

As with other Percoidei fishes, these fishes are characterized by the presence of spines in the dorsal, pelvic, and anal fins; the presence of two dorsal fins but no adipose fin; the abdominal placement of pelvic fins; pectoral fins that are placed laterally and oriented vertically; the presence of 17 or less principal rays in the caudal fin; and, usually, ctenoid scales (some species possess cycloid scales). These fishes are further characterized by a maxilla that is excluded from the gape of the jaw, a gas bladder that is physoclistous (lacking a connection between the gas bladder and the gut), and bone that is acellular. They also lack four specific bone types: the orbitosphenoid (within the orbital region of the skull), the mesocoracoid (within the pectoral girdle; this bone helps to position the pectoral fin obliquely rather than vertically, as in the percoids), the epileural (riblike intermuscular bones extending below the vertebral column), and the epicentral (primitively ligamentous, riblike intermuscular bones). Body shapes are quite divergent, and range from deep-bodied and highly compressed (Kyphosidae, Drepaneidae, Monodactylidae, Pentacerotidae, Chaetodontidae, Pomacanthidae, and Cheilodactylidae) to elongated (Mullidae) forms. Similarly, there is considerable

variation in the ecomorphology of the head, jaws, or other structures, both between and within families. Some families, such as the Chaetodontidae, Pomacanthidae, and Cirrhitidae, include many species with brightly colored or highly conspicuous color patterns. Other families (Nandidae) are noted for the cryptic coloration of many species.

Goatfishes are characterized by pairs of long, chemosensory barbels under the lower jaw, a terminal mouth, and the presence of large ctenoid scales. Goatfish larvae are laterally compressed and moderate in depth or elongate. The barbels begin to form at about a length of 0.3–0.35 in (0.8–0.9 cm). Archerfishes have moderately compressed bodies, a pointed head, a dorsal that originates well back towards the caudal fin, and a grooved palate that can direct a stream of water into the air. Their larvae are moderate in depth initially, but become deeper, compressed, and heavily pigmented with age. Galjoens have deep bodies, small ctenoid scales (even on the fins), and a scaleless snout. Sea chubs have a small head with a terminal mouth, a deep and moderately compressed body, a dorsal fin that is continuous, and a caudal fin that is either emarginate (notched) or forked. Their larvae are initially elongate, but become moderate in depth with age, with moderate-to-heavy pigmentation. There is some variation between subfamilies. The jutjaw has an oval-shaped, compressed body, small ctenoid scales, and a large mouth with a projecting lower jaw. Sicklefishes have strongly compressed, oval-shaped bodies covered with cycloid scales and protusile mouths. Monos have deep, highly compressed bodies that are covered with small deciduous scales. Pelvic fins are present in juveniles, but become rudimentary or absent in adults. Their larvae are specialized, having head spination that is moderately developed and elongate pelvic fins that form rather early.

The butterflyfishes have deep, highly compressed bodies; small protractile mouths; somewhat small ctenoid scales; a single dorsal fin that is continuous; and a caudal fin that is rounded or slightly emarginate. The post-larvae have a distinctive tholichthys stage, denoted by the presence of bony plates and protruding spines on the head. Angelfishes also have deep, compressed bodies, along with small ctenoid scales, small mouths, and a single dorsal fin that is unnotched. Characters that differentiate angelfishes from the butterflyfishes are a single prominent spine located at the corner of the preopercle; smaller spines on the preopercle, interopercle, and preorbital; and the absence of a tholichthys postlarval stage. The oldwife, as an adult, has remarkably large dorsal, pelvic, and anal fins, as well as small ctenoid scales, and a deep, compressed body. Sharp spines are present on the head, anal fin, and dorsal fin; the dorsal spines contain poison and thus the oldwife must be handled with care. Larvae are elongate to moderately long, and are heavily pigmented.

Boarfishes and armorheads resemble the adult oldwife. They have deep, moderately to strongly compressed bodies, exposed head bones that form rough, striated plates, and may have elongated mouths and fin rays. Juveniles may have tall dorsal fins and elongated anal and pelvic fins, but these are reduced as adults. Some species have poisonous spines and should be handled with care. Their larvae are denoted by elongate pelvic fins and extensive, well-developed spination on the head. Leaffishes possess large heads and mouths (the mouth is protactile), continuous dorsal fins, and rounded caudal fins. Knifejaws (Oplegnathidae) have oblong, moderately compressed bodies, tiny ctenoid scales, a single dorsal fin, and a parrotlike beak reminiscent of that on the distantly related parrotfishes (Scaridae). Knifejaw larvae lack subopercular or interopercular spines and are lightly pigmented. Hawkfishes resemble small groupers and their allies (Serranidae) and are characterized by the presence of filaments arranged in small tufts on the tips of the dorsal fin spines and by lower pectoral fin rays that are thickened. Members of the genus Paracirrhites, especially P. arcatus, P. hemistictus, and P. forsteri, are polychromatic. Two fixed color morphs occur in the former two species, while the latter exhibits two fixed color patterns in juveniles and a wide range of patterns that have no apparent relationship to sex, adult body size, or geographic locality. However, geographical color variation has been reported for the wide-ranging Cirrhitus pinnulatus. Hawkfish larvae have a pigmented chip barbel that disappears with age, cirri on the tips of the dorsal fin spine membrane, and serrations on the preopercle.

Kelpfishes are grouper or hawkfish-like in appearance, with cycloid scales that are moderately large, large pectoral fins, filaments on the tips of dorsal fins, and tufts of filaments on the dorsal spines of some species. Larvae are elongate to moderate in length, lack head spines, have heavy pigmentation in the postflexion stage, and are unspecialized. Sea carps have robust, elongate bodies, small cycloid scales, and pectoral fins with thickened lower rays positioned well forward under the gill covers. Their larvae are long to very elongate, lacking in head spines, heavily pigmented in the postflexion stage, and rather unspecialized. Many morwongs have elongated pectoral rays, forked caudal fins, an elevated dorsal fin, and large rubbery lips. A humplike head and back is present in a number of species. The postlarval stage of morwongs is silvery in color, deep-bodied, and very thin. This stage is retained in most species until they reach 17.7–23.6 in (45–60 cm) in total length. Although somewhat similar in shape to morwongs, the trumpeters have bodies that are elongate and compressed, with small pectoral fins, dorsal fins that are deeply notched, and a protruding mouth. Their postlarval stage resembles that of the morwongs.

Distribution

These fishes occur in coastal and estuarine waters of the Atlantic, Pacific, and Indian Oceans. Some species also occur in freshwater habitats in Africa, Australia, South America, and Southeast Asia. The Mullidae (six genera and 55 species) is found in tropical and warm-temperate coastal waters of the Atlantic, Pacific, and Indian Oceans. The Toxotidae (one genus and six species) is distributed in coastal marine, brackish, and fresh waters of India, east to the Philippines and Vanuatu, and south to northern Australia. The Dichistiidae (one genus and three species) has a distribution limited to coastal and brackish waters in South Africa and Madagascar. The Kyphosidae (15 genera and 42 species) is mostly tropical or subtropical in distribution, but two species have adapted to temperate waters off California. The Paracorpididae (one genus and at least one species) is known from Mozambique and South Africa. The Drepaneidae (one genus and three species) ranges from the Indo-West Pacific to West Africa. The Monodactylidae (two genera and five species) includes popular aquarium fishes found in tropical and warm temperate waters. One wide-ranging species, Monodactylus argenteus, occurs in brackish, marine, and freshwater habitats from the Red Sea and East Africa, east to Samoa, north to the Yaeyama Islands of southern Japan, and south to Australia and New Caledonia. A second species, M. falciformis, is limited to the western Indian Ocean, from the Red Sea, down to South Africa, and another, M. kottelati, is found only at Sri Lanka. Monodactylus sebae is found in the eastern Atlantic, from the Canary Islands and Senegal south to Angola. The two remaining species, both in the genus Schuettea, are found in temperate brackish and marine waters of southern Australia; one species off southeastern Australia, and the other off Western Australia. The Chaetodontidae (10 genera and as many as 125 species) and the Pomacanthidae (nine genera and at least 73 species) are distributed in tropical, subtropical, and warm temperate coastal waters. Among the chaetodontids, at least 13 species occur in the Atlantic, four species in the eastern Pacific, and the remainder in the Indo-West Pacific region. A similar pattern exists for the pomacanthids. The Enoplosidae

(one genus and species) is found in coastal temperate waters of southern Australia. The Pentacerotidae (eight genera and 13 species) occurs primarily in temperate waters of the Indo-Pacific and southwestern Atlantic. The Nandidae (seven genera and 10 species) contains primarily freshwater fishes, but may also occur in brackish-water estuaries. Two species occur in riverine freshwaters of tropical western Africa. Polycentrus schomburgkii is found in fresh and brackish water from Trinidad and the Guyanas south to the Amazon Basin, while Monocirrhus polyacanthus is found from Guyana south to Brazil and west to the Peruvian Amazon. The remaining species occur from southern Asia east to Southeast Asia, including Indonesia. The Oplegnathidae (one genus and six species) is found in the coastal warm-temperate waters of the Indo-Pacific region, mainly in Japan, southern Australia (including Tasmania), the Galápagos Islands, Peru, and South Africa. One species has also been reported from the volcanic far-northern Mariana Islands. Among cirrhitoid fishes, the Cirrhitidae (12 genera and 33 species) occurs mainly in the Indo-West Pacific region, but also in the tropical and subtropical Atlantic (three species), and eastern Pacific (three species). Two species have distributions that range from the Red Sea east to Mexico and south to Colombia. Three other species range from the Red Sea east to Hawaii and eastern French Polynesia. On the other hand, a number of hawkfishes have limited distributions. For example, the giant hawkfish (Cirrhitus rivulatus) is restricted to the Eastern Pacific, including the Galápagos. Similarly, Itycirrhitus wilhelmi is found only at Easter and Pitcairn

Islands at the eastern margin of the central Pacific. The splendid hawkfish (Notocirrhitus splendens) is endemic to Lord Howe Island, but strays west to Sydney and the coast of New South Wales in Australia, but also occurs at the Kermadec Islands of New Zealand. The red-barred hawkfish (Cirrhitops fasciatus) has a disjunct distribution and is found only in Hawaii, southern Japan, Mauritius, and Madagascar. Another species with a disjunct distribution, Paracirrhites hemistictus, occurs sporadically throughout the central and western Pacific, but is also found at Christmas and Cocos-Keeling Islands in the eastern Indian Ocean. The Chironemidae (two genera and four species) is distributed in temperate coastal waters of Australia and New Zealand, with one species endemic to Chile, and another limited to the warm-temperate Lord Howe, Norfolk, and the Kermadec Islands. The Aplodactylidae (three genera and five species) is another Southern Hemisphere family found in the temperate marine waters of Australia, New Zealand, Peru, and Chile. The Cheilodactylidae (four genera and 20 species; Goniistius of some authors, treated here as a subgenus of Cheilodactylus) has an antitropical distribution. Most species are found in Southern Hemisphere waters (Australia, New Zealand; the Kermadec, Lord Howe, Norfolk, Easter, Rapa, St. Paul's, and Amsterdam Islands; Ilots de Bass, southern Africa, Chile and Peru in the Indo-Pacific; Argentina, southern Africa, Tristan da Cunta Island and Vima Mount in the Atlantic) but members of one genus, Cheilodactylus, also occur off Japan, China, and the Hawaiian Islands. Finally, another Southern Hemisphere family, the Latridae (three genera and nine species), is distributed in coastal southern Australia, New Zealand, Chile, and the southern Atlantic.

Habitat

The habitats of these fishes are highly variable, and range from coastal and deepslope or seamount marine habitats, to estuaries, rivers, lakes, ponds, swamps, and even water-filled ditches. Three terms describe the use of these habitats. Benthic fishes are those found in close association with the bottom (including structure); benthopelagic fishes swim on or just above the bottom; and pelagic fishes swim in the water column well above the bottom. Goatfishes are benthopelagic fishes that frequent coral reefs, rocky reefs, and sand, rubble, and mud flats. Archerfishes are pelagic and swim frequently just beneath the surface of shallow marine and brackish waters, and in freshwater lakes and streams. They prefer coastal and estuarine mangroves or weed beds, and flooded trees in freshwaters. Three species are restricted to freshwater habitats. Galjoens are pelagic shoaling fishes found in turbid water along rocky coastlines. Sea chubs are pelagic or benthopelagic, although some pelagic species may be benthopelagic at night. They occur on coral and rocky reefs and frequent high-energy or surge zones. Jutjaws are pelagic fishes of coastal waters. Sicklefishes are benthopelagic in coastal and estuarine waters, and frequent mudflats. Monos are pelagic or benthopelagic, and may frequent mangroves or other structures when not swimming in the water column of bays, back bays, estuaries, or the lower reaches of rivers. Butterflyfishes are either benthopelagic or pelagic over coral reefs (most species), rocky reefs, walls and deep slopes (pelagic species, such as Hemitaurichthys polycanthus, that feed on zooplankton), and flats. Juveniles and adults of some species have been reported from mangroves. Similarly, the angelfishes are either benthopelagic or semipelagic, the latter (i.e., Genicanthus spp.) moving up into the water column to feed upon zooplankton. A number of benthopelagic angelfishes, such as some members of the genus Centropyge, are often found among coral rubble, dead corals, or rocks, where filamentous algae, a component of their diet, is found. Adults of the single species of oldwife (Enoplosus armatus) are found on inshore and offshore rocky reefs, and on sea grass beds and in estuaries, but juveniles occur mainly in estuaries. The boarfishes and armorheads are pelagic and occur on deep slopes of inshore reefs, often in or near caves, and seamounts offshore. The leaffishes occur largely in freshwater. All are benthopelagic. Three riverine species are also found in brackish water in estuaries. The most famous leaffish, the Badis badis, lives solitarily in rivers, ponds, ditches, and swamps. All leaffishes make use of structure, usually plant materials, for shelter and ambush sites. The knifejaws are found swimming among rocks, underwater cliffs, and walls on rocky reefs. The hawkfishes are benthic, although one species, Cyprinocirrhites polyactis, hovers in the

water column to feed on zooplankton, but darts into holes on the bottom to avoid danger or to shelter for the night. Two species of hawkfishes, Neocirrhites armatus and Oxycirrhites typus, are obligate coral-dwelling fishes. The former species occurs in shrublike corals of the genus Pocillopora, while the latter occurs in gorgonians (sea fans) and black corals found on deep slopes and walls of coral and rocky reefs. Species of the genus Paracirrhites are faculative coral-dwelling fishes that shelter in or perch on larger shrub-like corals. If such corals are absent, they will also perch upon coral boulders, ledges and rocks, or hide in holes. Kelpfishes are bottom dwelling, usually in inshore kelp forests, algal beds, and surge zones, or in intertidal reaches of rocky reefs. Seacarps are bottom dwelling and poor swimmers. They are found in small or large loosely maintained aggregations, in surge zones of rocky reefs, and in tidal passes in estuaries. The morwongs occur in a variety of habitats and depth ranges; most species are found on coral or rocky reefs inshore to about 197 ft (60 m), but Nemadactylus species have been reported from deep-slope habitats down to 1,312 ft (400 m). The trumpeters occur on rocky reefs and bottoms, although members of the genus Mendosoma forage for zooplankton in the water column.

Behavior

There is considerable variety in social organization, locomotion, and other behavior among members of this group. General patterns characteristic of each family are presented here with some greater detail given to certain families. Goatfishes move purposefully along the bottom while foraging and may accompany other bottom-feeding or swimming species for opportunistic feeding, and other fishes, such as wrasses (Labridae), may join them for the same reason. Many goatfishes aggregate in small groups, although members of certain genera, such as Mulloidichthys form large schools that seem to hover in the water column when not foraging on the bottom. Others move about singly, however, and some of these, such as Parupeneus cyclostomus (a species with two color morphs, golden yellow and pale blue), are remarkably active swimmers. Many species also rest upon the bottom for long periods of time, while some are more active at night than during daytime. The archerfishes use stealth when hunting prey in the manner discussed, but they may be territorial as they move back and forth encountering their neighbors. Galjoens move about in small groups, as do the sea chubs, although the latter can form exceptionally large groups that move about in the water column. The behavior of the jutjaws is poorly known, but they likely form aggregations or schools that swim in the water column. Sicklefishes do the same closer to the bottom. Monos form large schools in the mouths of rivers or in bays, but some occur singly or in small groups under the cover of mangroves in brackish water or brush in freshwater, where some species may be highly territorial.

The behavior of butterflyfishes, perhaps because of their bright coloration and high level of conspicuousness, has been the subject of considerable research. Largely diurnal, butterflyfishes may occur singly and be highly territorial, or live in aggregations with a general home range and relatively little territorial or aggressive behavior. Most, however, may be found in pairs, usually monogamous, that patrol home ranges or protect shared territories. Territorial defense, in general, may be directed towards conspecifics, other butterflyfishes with similar feeding requirements or preferences, or other species of fishes that also share their diet. Species that live singly or in pairs may form aggregations during certain times of the day or season, and these may be related to reproduction. At least one species, Chaetodon lunula, which may form aggregations during daytime, is active at night.

The angelfishes mostly form social and mating groups consisting of a single male and multiple females. They change sex from female to male (protogynous hermaphroditism), usually under social control. Some species may be monogamous or facultatively monogamous. For example, the emperor angelfish (Pomacanthus imperator) may often be encountered in pairs that consist of a male and female, yet additional females may reside in the pair's home range and join them prior to courtship at dusk. The regal angelfish (Pygoplites diacanthus) is often seen alone, but will form pairs or mating groups during sunset courtship periods. Some intra- and interspecific territorial behavior may occur, usually during encounters while patrolling a home range.

The behavior of the oldwife is not well known. Juveniles occur singly, but tend to form schools as they age. Adults are encountered either singly or in pairs, but large schools may also form. Shallow-dwelling boarfishes occur in pairs or small-to-large aggregations, depending upon the species, but the rarely seen juveniles occur singly. The behavior of their deep-water cousins, the armorheads, is poorly known, but they likely aggregate or form schools. The leaffishes are well known for the way they mimic leaves drifting in the water column. This allows them to avoid predation and to ambush potential prey. Little is known about the behavior of knifejaws except that they swim about rocks and will enter caves.

The hawkfishes are territorial and organized socially into mating groups consisting of a single male and a variable number of females (usually two to five) that maintain territories or home areas inside the male's territory. Two species, however, are facultatively monogamous as a consequence of habitat use. For example, obligate coral-dwelling species, such as Oxycirrhites typus and Neocirrhites armatus, are facultatively monogamous if their coral (black coral or gorgonian sea fan, in the case of O. typus) is too small to support more than two fish, and if neighboring corals are too far away to allow a male to safely move to it to court a resident female. However, if the coral is large enough to support more than two fishes, or if suitable corals are close by, then a multiple-female group is possible. Juvenile obligate coral-dwelling hawkfishes often recruit to less-favorable corals nearby and then attempt, as do facultative and noncoral-dwelling species, to gain entry into the mating group. Usually, resident females will attack the new arrival and try to expel it from the coral or the group. This behavior is also directed towards adults, usually emigrants from fragmented groups elsewhere, that attempt to join the group. If the emigrant is recognized as another male, the resident male will attempt to expel it; if a female, it may be attacked by other females. As with other fishes that have a similar social organization, all hawkfishes likely change sex from female to male (protogynous hermaphroditism). Sex change may proceed through social control or maturation. The effect of sex change upon social organization can be profound, because, in addition to allowing for succession by the dominant female with the loss of the male, it can also allow the dominant female to fragment the group should it become too large for the male to control effectively. Similarly, if the dominant male cannot control his group, then neighboring males and "rogue" males (those that have lost a group or were recent females that could not fragment their own group after changing sex) can attempt to sequester females. Another interesting behavior is practiced by the plankton-feeding Cyprinocirrhites polyactis, which mimics fairy basslets (Serranidae: Anthiinae) that aggregate, often in considerable numbers, in the water column.

Kelpfishes may establish large aggregations that form loosely over reefs, particularly surge zones, but individuals and small groups are more likely in the intertidal zone. The seacarps also aggregate, either in small or large, loosely organized groups that rest upon the bottom. They swim with the dorsal fin raised when seeking food or changing location. Most morwongs move about the bottom singly or in small, loosely formed aggregations, but members of the genus Nemadactylus form large aggregations. Trumpeters are mostly schooling fishes, which move about in small groups or in aggregations of hundreds of individuals.

Feeding ecology and diet

These fishes have considerable variation in both feeding methods and diet. Although most direct their attention towards bottom feeding or planktivory, in which they feed upon phytoplankton or zooplankton in the water column, there are some remarkable exceptions. Goatfishes utilize their barbels, which bear chemosensory organs, to stir up the bottom and

detect prey. They also feed upon smaller invertebrates in sand, rubble, algal beds, or on coral pavement, and some species may take small fishes that they disturb. Archerfishes swim near the surface of the water and can detect visually, with a correction for light refraction, terrestrial prey on overhanging branches of trees and shrubs or blades of grass at the water's edge. Upon detecting the prey, usually an insect, archerfishes direct a stream of water drops toward that prey, knocking it to the water where it can be captured with the mouth and consumed. The drops are formed by compression of water by the gill covers and shot by forcing them along a groove formed by the tongue and palate. Archerfishes also consume floating fruits and flowers. Although galjoens have small mouths, they are capable of preying upon ascidians, mussels, barnacles, and other crustaceans. Galjoens also consume seaweeds. The sea chubs have a mixed feeding strategy and diet. Members of the subfamilies Kyphosinae (except the genus Gras) and Girellinae graze or pluck algae. Those in the subfamilies Scorpidinae and Microcanthinae are carnivores that feed upon benthic invertebrates. The jutjaws strain zooplankton from the water column. The sicklefishes feed upon small invertebrates they extract from soft sediments, usually mud. Monos have small, obliquely positioned mouths with a combination of brushlike teeth on their jaws, villiform teeth on the vomer, and palatines within the mouth. They use this combination to capture and feed upon small fishes, invertebrates, or plankton. The butterflyfishes have been collectively described as microconsumers. Some species feed exclusively upon coral polyps, others upon small invertebrates on corals, rocks, coral pavement, or crevices, and still others, such as Hemitaurichthys polylepis, forage for zooplankton in the water column. There is some plasticity in diet for a number of species. While Chaetodon trifascialis feeds only upon acroporid corals, C. punctatofasciatus will feed upon a variety of benthic invertebrates, corals, and even filamentous algae. The forcepsfish (Forcipiger flavissimus) uses its tubular snout to feed upon hydroids, fish eggs, and small crustaceans, and to excise pieces of the tentacles of polychaete worms, the tube feet of starfishes, and the pedicilaria of sea urchins. On the other hand, the longnose butterflyfish (F. longirostris), with its longer snout, feeds mainly upon small crustaceans that are taken whole. Angelfishes also

have considerable plasticity in their diet, although perhaps less so compared to butterflyfishes. Pygmy angelfishes (Centropyge spp.) feed mostly upon filamentous algae, while Genicanthus angelfishes forage mainly for zooplankton in the water column and supplement their diet with benthic invertebrates or algae. Others in the family feed upon sponges, small soft-bodied invertebrates, fish eggs, and algae.

Little is known about the feeding habits and diets of the oldwife, boarfishes, and armorheads, but they appear to feed upon benthic invertebrates and fishes. Leaffishes, which can conceal themselves even in open water thanks to their ability to mimic leaves downed in water, are remarkably good predators upon fishes and invertebrates (mostly crustaceans, insects, and worms). However, one larger species from Southeast Asia, the catopra (Pristolepis fasciata), is primarily a herbivore that feeds upon filamentous algae, submerged land plants, seeds, and fruits, but will also take crustaceans and aquatic insects. Knifejaws use their parrotlike beaks to break open and feed upon barnacles and mollusks. The hawkfishes feed upon small crustaceans and fishes they ambush from a perched position on a coral or rock, or from a resting position on the bottom. One species, Cyprinocirrhites polyactis, plucks zooplankton while hovering in the water column, and a second species, Cirrhitichthys aprinus, has been observed taking zooplankton as it darts out from soft corals into the water column along current-swept reef walls. Kelpfishes exhibit feeding behavior similar to hawkfishes and feed upon small invertebrates. The seacarps feed upon algae and other seaweeds they grab from the bottom with their mouths. The small mouths and thick lips of morwongs are used to feed upon small benthic or pelagic invertebrates, such as polychaete worms, crustaceans, mollusks, and echinoderms, or in some species, algae. At least one species, Chirodactylus grandis, feeds upon squid and small fishes. Trumpeters feed upon benthic invertebrates, although the members of the genus Mendosoma feed upon zooplankton.

Reproductive biology

Although studies on reproductive biology of these fishes are far from complete, there is considerable information, which characterizes some families in detail. Overall, they are either gonochorists or sequential hermaphrodites. Gonochoristic fishes begin life either as females or males and remain that way their entire lives. Examples include goatfishes, archerfishes, galjoens, sea chubs, jutjaws, sicklefishes, monos, butterflyfishes, the oldwife, boarfishes and armorheads, leaffishes, knifejaws, and morwongs. Sequential hermaphrodites change sex. Most Percoidei with this life strategy appear to be protogynous hermaphrodites, in that they begin as females but may change sex and become males. The control of sex change may be social or as a consequence of maturation, and despite detailed research on some groups (i.e., angelfishes and hawkfishes), much remains to be learned about the circumstances that trigger sex change. The status of kelpfishes, seacarps, and trumpeters is uncertain.

Spawning may occur in monogamous pairs, as sequential pairs within a haremic mating group or spawning aggregation, or promiscuously. Most of these fishes appear to spawn pelagically, the eggs are fertilized in the water column during pair spawning, and there is no parental care. Sneaking (partial fertilization of a pair's eggs by a second male that sneaks up on a rising pair) and group spawning are also possible. Pelagic spawning has been verified for the goatfishes (paired and group spawning), archerfishes, sea chubs, monos (in part), butterflyfishes, angelfishes, the oldwife, hawkfishes, morwongs, and trumpeters. Pelagic spawning is presumed in the galjoens, jutjaws, sicklefishes, boarfishes and armorheads, knifejaws, kelpfishes, and seacarps. Demersal spawning, in which eggs are scattered on the bottom, placed in nests or bubble nests (a specialized behavior of some leaffishes), brooded in holes or caves, or attached to plants, shells, rocks, or some form of structure, may involve parental care to some degree or not at all. Parental care is pronounced, especially in freshwater fishes such as the leaffishes, but is largely absent or minimal in marine fishes. Leaffishes spawn demersally, or in bubble nests, and some monos also have demersal eggs.

There is considerable variability in the temporal patterns of spawning of these fishes. They may spawn only seasonally in relation to latitude and water temperature but not exclusively.

For example, fishes of some families might spawn seasonally over a short period of time once or twice a year regardless of latitude. Others, such as the hawkfishes, may spawn daily at lower latitudes, but seasonally at higher latitudes. Those fishes that produce pelagic eggs and larvae have the potential for relatively long-distance dispersal in marine systems. Demersally spawned eggs usually also hatch to become pelagic larvae, but these may not disperse as far. Of course, there are plenty of exceptions to the rule in both cases.

Conservation status

One species of angelfish (Centropyge resplendens [Pomacanthidae]), and five species of butterflyfish (Chaetodon flavocoronatus, C. litus, C. marleyi, C. obliquus, and C. robustus [Chaetodontidae]) are listed as Vulnerable on the IUCN Red List. Other species may be vulnerable or threatened, but the data simply do not exist to verify the conservation status of many of these fishes. Certainly, species that live in close association with corals (such as butterflyfishes, angelfishes, and hawkfishes) might be at risk because of the effects of coral bleaching, the harvest of corals, and the general destruction of coral habitats. The same might be said of species that live in mangroves (such as archerfishes) threatened with destruction. The leaffishes may be at risk from the effects of pollution, hydroelectric development, so-called stream improvements, the draining of swamps, destructive agricultural practices, mining, logging, and other forms of development that contribute to the

sedimentation of rivers and streams. Galjoens require considerable protection from overfishing in South Africa. Similarly, armorheads that dwell on seamounts or on deep-slope reefs and grounds are likely candidates for overfishing and should be of considerable concern. Overfishing of popular aquarium species (such as leaffishes, butterflyfishes, angelfishes, and hawkfishes), especially those that are rare, have limited distributions locally or geographically, or have mating systems or other life history characteristics that make them especially vulnerable to unregulated harvests, is also of great concern.

Significance to humans

Fishes of the following families are taken in major or minor commercial food fisheries: Mullidae, Parascorpididae, Enoplosidae, Pentacerotidae, Oplegnathidae, Cheilodactylidae, Chironemidae, and Latridae. Fishes that may be found in subsistence food fisheries include the Mullidae, Toxotidae, Kyphosidae (prized at some localities but avoided at others), Monodactylidae, Chaetodontidae, Pomacanthidae, Nandidae, larger Cirrhitidae, and Cheilodactylidae. Game fishes include members of the Dichistiidae (their numbers heavily depleted in South Africa), Nandidae, Oplegnathidae, Cheilodactylidae, and the Latridae. Some members of the Chironemidae and Aplodactylidae are not taken generally for food because they apparently taste bad. A number of families are taken for the aquarium trade, either as juveniles or adults. Important species include members of the Toxotidae, Nandidae, Monodactylidae, Chaetodontidae, Pomacanthidae, and Cirrhitidae. Of minor importance are members of the Mullidae, Kyphosidae, Enoplosidae, and Cheilodactylidae.

Species accounts

List of Species

Pebbled butterflyfish

Chaetodon multicinctus

family

Chaetodontidae

taxonomy

Chaetodon multicinctus Garrett, 1863, Hawaiian Islands.

other common names

Hawaiian: Kikakapu.

physical characteristics

Deep, compressed body, whitish in color, with four to six narrow brown or gold-brown bars and many faint olive spots on both the body and the fins; a gold-brown eye bar on the head, a black and gold bar on the caudal peduncle, and a solid black bar on the caudal fin. To 4.5 in (12 cm) total length.

distribution

Limited to the Hawaiian Islands and Johnston Atoll in the east-central Pacific.

habitat

Seaward or lagoon coral reefs, usually with considerable stands of Porites and Pocillopora corals, between 16 and 98 ft (5–30 m) in depth.

behavior

Forms heterosexual pairs but occasionally occurs in small aggregations.

feeding ecology and diet

Omnivorous, its diet consisting of coral polyps, polychaete worms, small crustaceans, and algae.

reproductive biology

Gonochoristic. Paired courtship just before or after sunset into early evening, with pelagic spawning in the water column. Spawning pairs are occasionally joined by one or more intruding males who attempt to spawn with the female. The intruders may be either rogue males or paired males who have temporarily abandoned their mates. Courtship occurs between December and July, but courtship and spawning is most pronounced between March and July. Eggs and larvae are pelagic. The eggs are spherical and small (0.023–0.029 in [0.6–0.75 mm] in diameter). Larvae are around 0.059 in (1.5 mm) in length at hatching and have a large yolk sac. The mouth is unformed and the eyes are unpigmented. With growth, the larva develops fused head plates that extend over the trunk of the body. This is known as the thoichthys stage of development; it adapts the larva to a long pelagic phase. Other adaptations include spine formation on the head, dorsal, and pelvic fins.

conservation status

Not listed by the IUCN. Coral habitats may be threatened by coral bleaching, pollution, sedimentation, and other forms of degradation that may negatively impact populations of this limited-distribution species.

significance to humans

May be collected for the aquarium trade, but this and many other butterflyfishes generally do poorly in most aquaria.

Hawaiian morwong

Cheilodactylus vittatus

family

Cheilodactylidae

taxonomy

Cheilodactylus vittatus Garrett, 1864, Hawaiian Islands.

other common names

Hawaiian: Kikakapu.

physical characteristics

Laterally compressed with a high forehead and long, sloping back. There are prominent bony knobs on the head. The caudal fin is forked. There is an oblique pale black bar on the head, and there are four oblique dark-black bars, all against a white background color, along the head and flank. The dorsal fin spines are black, white, and orange-red; the dorsal rays are white.

distribution

Hawaiian Islands in the Northern Hemisphere, and New Caledonia, Lord Howe Island, and the Kermedec Islands in the Southern Hemisphere.

habitat

Coral and volcanic rocky reefs, in areas of coral, rocks, pavement and rubble. Depth usually below 65 ft (20 m).

behavior

Occur singly or in small groups or aggregations.

feeding ecology and diet

Benthic carnivores that forage on the bottom for invertebrates, mainly crabs and shrimps, amphipods, polychaete worms, and gastropod mollusks.

reproductive biology

Gonochoristic, with females tending to be larger than males. Courtship and spawning at night, with the release of pelagic eggs and larvae. Eggs are spherical and range in size from about 0.035 to 0.043 in (0.9–1.1 mm) in diameter. Hatching larvae are approximately 0.1–0.13 in (2.5–3.3 mm) in length, have unpigmented eyes, an unformed mouth, and a large yolk sac. Pigmentation develops with absorption of the yolk. The body is elongate but becomes deeper and compressed with growth, and silvery in color (paper-fish phase). The ventral keel also becomes prominent with growth. The gas bladder is small to moderate in size. The mouth becomes smaller and oblique with development. Larvae are adapted for a relatively long pelagic life.

conservation status

Not listed by the IUCN. This species is rare in the Hawaiian Islands and may yet be proven distinct from those of the Southern Hemisphere populations.

significance to humans

Taken as a food fish, but smaller individuals may also be collected for the aquarium trade.

Longnose hawkfish

Oxycirrhites typus

family

Cirrhitidae

taxonomy

Oxycirrhites typus Bleeker, 1857, Ambon Island, Moluccas, Indonesia.

other common names

German: Gestreifter Schützenfish; Japanese: Kudagonbe.

physical characteristics

Body slender, with an elongated snout; body whitish in color with a tartanlike pattern in red along body, snout, and dorsal fin; males larger than females within mating groups. To 5.1 in (13 cm).

distribution

Red Sea east to Panama; southern Japan south to New Caledonia.

habitat

Usually deep slope or wall habitats in seaward coral or rocky reefs, passes, and lagoons; shallow rocky reefs in southern Baja California Sur, Baja Peninsula, Mexico, and occasionally found on shallow coral reefs elsewhere, particularly in areas of upwelling. Otherwise, more common in deeper reef habitats. Usually found in gorgonians (sea fans) and black corals.

behavior

Occurs singly or in facultatively monogamous pairs on gorgonians and black corals. Will have a haremic mating system if the gorgonian or black coral is large or if others are close by.

feeding ecology and diet

Plucks small benthic and planktonic crustaceans with its long snout.

reproductive biology

Protogynous hermaphrodites. Paired courtship and pelagic spawning just prior to or after sunset. If a male has a haremic mating group, he will move back and forth between females to court them until they spawn. Eggs and larvae are pelagic. Eggs are spherical, about 0.027 in (0.69 mm) in diameter, and hatch after 15 hours incubation at 80.6°F (27°C). The yolk sac is

large, and the jaws and eyes undeveloped. Pigmentation develops with absorption of the yolk. Larvae are elongate and compressed laterally. Possess a gas bladder that is lost in adults. Specializations for pelagic life include serrations on the preopercle, a chin barbel (which disappears with growth), and the presence of cirri on the spinous dorsal fin membrane and also on the anterior nostril.

conservation status

Not listed by the IUCN. Not commonly seen, despite its wide distribution, and population levels likely to be limited by available microhabitat. The collection of black corals for the jewelry trade and gorgonians for the ornamental trade poses a threat, as do unregulated harvests by the aquarium trade.

significance to humans

A highly prized aquarium species.

Cortez chub

Kyphosus elegans

family

Kyphosidae

taxonomy

Kyphosus elegans Peters, 1869, Mazatlan, Sinaloa, Mexico.

other common names

Spanish: Chopas.

physical characteristics

Deep, compressed body with silver-gray color marked by faint brown stripes between scale rows on the flanks. Can quickly produce a pattern of white spots about the diameter of the eye. The jaw teeth are fixed and not freely movable. To 15 in (38 cm) total length.

distribution

Gulf of California south to Panama and west to the Galápagos Islands.

habitat

Rocky and coral reefs, usually in high-energy areas below the surf zone.

behavior

Often found in large schools swimming in the water column.

feeding ecology and diet

Omnivorous but feeds mainly upon benthic algae.

reproductive biology

The eggs are pelagic and about 0.039–0.043 in (1–1.1 mm) in diameter. Larvae hatch at 0.094–0.114 in (2.4–2.9 mm), are elongate, have a large yolk sac, unformed jaws, and unpigmented eyes. With growth, the head and mouth become large. Possesses a gas bladder. Head spines develop as a specialization for pelagic life.

conservation status

Not listed by the IUCN.

significance to humans

May be taken in subsistence fisheries or local commercial fisheries as a food fish.

Mono

Monodactylus argenteus

family

Monodactylidae

taxonomy

Monodactylus argenteus Linnaeus, 1758, East Indies.

other common names

English: Diamondfish, fingerfish, moony, silvery moony, Natal moony; Japanese: Himetsubameuo.

physical characteristics

Deeply compressed body with silvery color marked by a slender dark-colored bar on the head. The anterior edges of the dorsal and anal fins are also dark colored, the fins are pale yellow. The pelvic fin is absent in adults. To 8.5 in (22 cm) total length.

distribution

Red Sea east to Samoa; Yaeyama Islands (southern Japan), south to Australia and New Caledonia.

habitat

Brackish waters, estuaries, and silty reefs.

behavior

Found singly or in small groups among mangroves or other structures, but in large schools in estuaries and silty reefs where they swim in the water column.

feeding ecology and diet

Feed upon plankton in the water column and detritus on the bottom.

reproductive biology

Spawns adhesive, demersal eggs in freshwater. Others in the genus spawn pelagic eggs in seawater. Eggs are spherical and approximately 0.039 in (1 mm) in diameter. The larvae hatch around 0.071 in (1.8 mm) in length, are compressed and moderate in depth, have a large yolk sac, an unformed mouth, and unpigmented eyes. With growth, the head becomes large with a convex dorsal profile that becomes steep over time. Possesses a small gas bladder. Larvae have pelvic fins that are reduced in adults. This character, plus the development of spines on the head, are viewed as specializations for pelagic life.

conservation status

Not listed by the IUCN.

significance to humans

Popular as an aquarium fish. Maybe taken in subsistence fisheries as a food fish.

Yellow goatfish

Mulloidichthys martinicus

family

Mullidae

taxonomy

Mulloidichthys martinicus Cuvier and Valenciennes, 1829, Martinique Island, West Indies.

other common names

French: Poisson chèvre jaune.

physical characteristics

Twin barbels on the lower jaw, a relatively short snout, the body elongate with two well-separated dorsal fins, two spines in the anal fin, and a forked caudal fin. There are 36–37 lateral line scales from the upper gill opening to the base of the caudal fin. These fishes are bronze and pale yellow in color with a yellow median stripe that extends along the length of the body, and grow to 12 in (30 cm) total length.

distribution

Western Atlantic, from Bermuda southwest to Florida and the Gulf of Mexico, and south from the Bahamas to Brazil.

habitat

Coral reefs and flats of sand or rubble.

behavior

Swims lazily or hovers in schools above the bottom during the day and forages after dark.

feeding ecology and diet

Feeds by probing the bottom with its barbels in search of prey, usually at night or during low-light periods. The diet includes annelid worms, crabs, ophiurans, and to a lesser extent, small fishes.

reproductive biology

Spawns in aggregations and produces pelagic eggs and larvae. Eggs are spherical and small, between 0.024–0.036 in (0.63–0.93 mm) in diameter. Larvae hatch between 0.063–0.134 in (1.6–3.4 mm) in length, are elongate, and have a large yolk sac, unformed jaws, and unpigmented eyes. Body pigmentation develops with absorption of the yolk sac. Possesses a gas bladder and a short gut. The barbels form at about 0.67–0.71 in (17–18 mm) but remain recessed under the lower jaw. Post-larvae appear capable of a relatively long pelagic phase prior to settlement.

conservation status

Not listed by the IUCN.

significance to humans

Highly esteemed as a food fish and taken in subsistence and local commercial fisheries.

Amazon leaffish

Monocirrhus polyacanthus

family

Nandidae

taxonomy

Monocirrhus polyacanthus Heckel, 1840, Marabitanos, Brazil.

other common names

English: Barbeled leaf fish, South American leaf-fish; German: Blattfisch; Spanish: Pez hoja.

physical characteristics

Total length 3.9 in (10 cm). Body somewhat elongated, compressed, and basslike. Profile of forehead is steep and rises to the first dorsal spine at a point directly over the posterior edge of pelvic fin. A narrow, lurelike organ extends forward from the lower part of a protrusible mouth. Sharp dorsal, anal, and pelvic spines. Body color yellowish bronze to light or dark brown. Outer edges and tips of the anal, pelvic, and caudal fins are dark brown. Narrow dark brown stripe runs obliquely from a point midway up the forehead down through the eye, a second similar stripe runs obliquely from the lower operculum to the eye. A third, fainter band is found mid flank below the dorsal fin.

distribution

South America, from Guyana south to Brazil and west into the Peruvian Amazon.

habitat

Benthopelagic, primarily streams and rivers; often in association with or near submerged vegetation.

behavior

Cryptic. Often found floating motionless, like a leaf, or in the cover of aquatic vegetation.

feeding ecology and diet

Carnivorous. An ambush predator that preys upon smaller fishes, and aquatic insects and their larvae. Engulfs prey rapidly by extending its large, protrusible mouth.

reproductive biology

Gonochoristic. Demersal eggs are spawned and fertilized in a nest prepared by the male. Male parental care of eggs. Eggs hatch after three to four days and are free swimming by five days.

conservation status

Not listed by the IUCN.

significance to humans

Collected commercially for the aquarium trade.

Gangetic leaffish

Nandus nandus

family

Nandidae

taxonomy

Nandus nandus Hamilton, 1822, Gangetic Provinces of India.

other common names

None known.

physical characteristics

Typically basslike; pale olive-bronze color with loosely arranged vertical brownish olive bands along the flank and extending into dorsal fin. Noted for its sharp dorsal, pelvic, and anal spines.

distribution

From Pakistan east to Thailand.

habitat

Benthopelagic in freshwater ponds, ditches, flooded fields, lakes, reservoirs, canals, rivers, and estuaries.

behavior

Often cryptic and holding to cover from where it can ambush prey.

feeding ecology and diet

Gangetic leaffishes are predators that feed upon smaller fishes and aquatic insects.

reproductive biology

Gonochoristic. Males clean a spawning site, usually a rock or leaf, and then court the female, who spawns around 200–400 demersal eggs that are fertilized by the male. Male parental care has been observed in aquaria.

conservation status

Not listed by the IUCN.

significance to humans

A prized food fish. Juveniles may also be taken for the aquarium trade.

Striped boarfish

Evistias acutirostris

family

Pentacerotidae

taxonomy

Evistias acutirostris Temminck and Schlegel, 1844, Nagasaki, Japan.

other common names

English: Whiskered armorhead; Japanese: Tenguidai.

physical characteristics

Body deep with a sail-like dorsal fin; the lower jaw has many small barbels. Color pattern denoted by a dark brown face and alternating dark brown and white vertical bands along the

body; all fins are yellow except for the pelvic fins, which are dark brown. To 25.6 in (65 cm) total length.

distribution

Antitropical distribution, with populations in central and southern Japan, including the Izu and Ogasawara Islands, and Hawaii in the Northern Hemisphere and Lord Howe Island (Australia), and the islands of northern New Zealand in the Southern Hemisphere.

habitat

Generally a deep-slope fish, from 65 to almost 1,000 ft (20 to almost 330 m) deep, depending upon locality. Found among rocky and coral reefs, usually in or near caves.

behavior

May be found in pairs or small groups.

feeding ecology and diet

Uses its snout and barbels to probe the bottom for various invertebrates.

reproductive biology

Gonochoristic. Presumed to spawn pelagically, but little is known of its reproductive behavior and ecology. Eggs and larvae are pelagic.

conservation status

Not listed by the IUCN. Might be vulnerable because of its status as a commercial food fish.

significance to humans

Esteemed as a food fish, but juveniles and young adults are also taken, although infrequently, for the aquarium trade.

Emperor angelfish

Pomacanthus imperator

family

Pomacanthidae

taxonomy

Pomacanthus imperator Bloch, 1787, Japan.

other common names

Afrikaans: Keiser-engelvis; French (Polynesia): Poisson empereur; paraharaha; Japanese: Tatejima-kinchakudai.

physical characteristics

Deeply compressed, with longish pelvic fins and a long preopercule spine. Adults and juveniles have distinct color phases. Adults are yellow with horizontal blue stripes along the flank, a deep purplish mask over the eye and on the flank above the pectoral fin, a white snout and mouth, and yellow fins with purplish blue stripes on the anal fin. Juveniles are deep navy blue to almost black, with concentric white circles on the flank and white bands on the head. One of the largest angelfishes, to over 15.7 in (40 cm) total length.

distribution

Red Sea east to the Line Islands and the Tuamotu Archipelago; southern Japan south to northern Australia, and southeast to Lord Howe Island and the Austral Islands. Reported from Hawaii, but either very rare as a stray or introduced from aquarium stock.

habitat

Coral reefs. Juveniles on patch reefs in lagoons, protected areas in lagoon passes, and among corals, in holes or under ledges on outer reefs or reef flats; subadults move to surge channels and holes on reef fronts; and adults are found usually near caves and ledges of seaward reefs, passes, or lagoons.

behavior

Usually observed paired or singly as they patrol a large home range; may display some intra- or interspecific territorial behavior (usually towards other similarly sized angelfishes). Have an haremic mating group, with subordinate females residing within the male's home range.

feeding ecology and diet

Feed almost exclusively on sponges and tunicates.

reproductive biology

Protogynous hermaphrodites. Sequential courtship and spawning among members of a mating group occur at dusk. Eggs and larvae are pelagic. Eggs are round, approximately 0.023–0.039 in (0.6–1 mm) in diameter. Larvae hatch around 0.05–0.1 in (1.3–2.6 mm) in length, have a large yolk sac, an unformed mouth, and unpigmented eyes. With growth, they assume a deep, laterally compressed shape, and the profile of the head becomes steep. Possesses a gas bladder. The development of head spination and spinules (spine-like structures on the skin) are specializations for pelagic life.

conservation status

Not listed by the IUCN. Because of apparent low population densities at many localities, this species is subject to overfishing by the aquarium trade.

significance to humans

Highly prized, especially the juveniles, as an aquarium fish. Adults are also taken occasionally in subsistence food fisheries.

Banded archerfish

Toxotes jaculatrix

family

Toxotidae

taxonomy

Toxotes jaculatrix Pallas, 1767, Indian Ocean.

other common names

None known.

physical characteristics

Body is compressed and almost triangular, with a pointed head. Color is silvery white with five black bands across the upper sides of the body, the dorsal fin, and the caudal peduncle, and yellow on anal, caudal, and dorsal fins. These fishes grow to 8 in (20 cm) total length.

distribution

Widespread from India and Sri Lanka, east through Southeast Asia, the Ryukyu Islands, Palau and Yap in Micronesia, Melanesia, and parts of Queensland in northern Australia; also in Vanuatu.

habitat

Generally found in bays, estuaries, and the lower reaches of rivers and creeks, usually among mangrove branches and roots.

behavior

Occurs singly or in small groups around shelter. Swims slowly as it hunts for prey both below and above the surface of the water. Will swim rapidly on occasion to challenge conspecifics. As with other archerfishes, the banded archerfish will hunt by shooting a stream of water drops at terrestrial prey above the surface.

reproductive biology

Gonochoristic, with pair spawning and the release of demersal eggs near the bottom. Others in the genus release either pelagic or demersal eggs. Demersal eggs are larger compared to pelagic eggs in this species, about 0.023–0.031 in (0.6–0.8 mm) in diameter. Larvae hatch at 0.12 in (3 mm) in length, are pelagic, possess a large yolk sac, have an unformed mouth, and lack pigmented eyes. With growth, the body becomes moderate in depth and the mouth large. Possesses a gas bladder. The development of head spines is regarded as a specialization for pelagic life.

conservation status

Not listed by the IUCN.

significance to humans

Taken in subsistence fisheries and as a light-tackle gamefish, but more importantly as an aquarium fish.

Resources

Books

Allen, G. R., and D. R. Robertson. Fishes of the Tropical Eastern Pacific. Honolulu: University of Hawaii Press, 1994.

Allen, G. R., and R. Swainston. The Marine Fishes of North-Western Australia. Perth: Western Australia Museum, 1988.

Allen, G. R., S. H. Midgley, and M. Allen. Field Guide to the Freshwater Fishes of Australia. Perth: Western Australian Museum, 2002.

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

Donaldson, T. J. "Assessing Phylogeny, Historical Ecology, and the Mating Systems of Hawkfishes (Cirrhitidae)." In Proceedings of the 5th Indo-Pacific Fish Conference, Noumea 1997, edited by B. Seret and J. Y. Sire. Paris: Societe Francaise d'Ichtyologie & Institut de Recherche pour le Developement, 1999.

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. Oxford: Blackwell Science, 1997.

Kuiter, R. H. Coastal Fishes of South-Eastern Australia. Honolulu: University of Hawaii Press, 1993.

Leis, J. M., and B. M. Carson-Ewart, eds. The Larvae of Indo-Pacific Coastal Fishes. Leiden: Brill, 2000.

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. 3rd ed. Barrigada, Guam: Coral Graphics, 1999.

Neira, F. J., A. G. Miskiewicz, and T. Trnski, eds. Larvae of Temperate Australian Fishes. Laboratory Guide for Larval Fish Identification. Perth: University of Western Australia Press, 1998.

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

Sadovy, Y., and A. S. Cornish. Reef Fishes of Hong Kong. Hong Kong: Hong Kong University Press, 2000.

Sadovy, Y., and A. C. J. Vincent. "Ecological Issues and the Trades in Live Reef Fishes." In Coral Reef Fishes: Dynamics and Diversity in a Complex Ecosystem, edited by P. F. Sale. San Diego: Academic Press, 2002.

Sakurai, A., Y. Sakamoto, and F. Mori. Aquarium Fish of the World. The Comprehensive Guide to 650 Species. San Francisco: Chronicle Books, 1993.

Schafer, F. Aqualog: Reference Fish of the World. All Labyrinths: Bettas, Gouramis, Snakeheads, Nandids. Morfelden-Walldorf, Germany: Verlag A. C. S. GmbH, 1997.

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

Thomson, D. A., L. T. Findley, and A. N. Kerstitch. Reef Fishes of the Sea of Cortez. 2nd ed. Tucson: University of Arizona Press, 1987.

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

Periodicals

Donaldson, T. J. "Facultative Monogamy in Obligate Coral-Dwelling Hawkfishes (Cirrhitidae)." Environmental Biology of Fishes 26 (1989): 295–302.

——. "Reproductive Behavior and Social Organization of Some Pacific Hawkfishes (Cirrhitidae)." Japanese Journal of Ichthyology 36 (1990): 439–458.

Donaldson, T. J., and P. L. Colin. "Pelagic Spawning of the Hawkfish Oxycirrhites typus." Environmental Biology of Fishes 24 (1989): 295–300.

Donaldson, T. J., J. T. Moyer, R. F. Myers, and P. J. Schuup. "Zoogeography of the Fishes of the Mariana, Ogasawar, and Izu Islands: A Preliminary Assessment." Natural History Research Special Issue 1 (1994): 303–332.

Lobel, P. S. "Spawning Behavior of Chaetodon multicinctus (Chaetodontidae); Pairs and Intruders." Environmental Biology of Fishes 25 (1989): 12–130.

Patterson, C., and G. D. Johnson. "The Intermuscular Bones and Ligaments of Teleostean Fishes." Smithsonian Contributions in Zoology 559 (1995): 1–83.

Randall, J. E. "Review of the Hawkfishes (Family Cirrhitidae)." Proceedings of the United States National Museum 114 (1963): 389–451.

——. "A Review of the Fishes of the Subgenus Goniistius, Genus Cheilodactylus, with Description of a New Species from Easter Island and Rapa." Occasional Papers of the Bernice P. Bishop Museum 25, no. 7 (1983): 1–24.

——. "Revision of the Generic Classification of the Hawkfishes (Cirrhitidae), with Descriptions of Three New Genera." Zootaxa 12 (2001): 1–12.

Sadovy, Y., and T. J. Donaldson. "Sexual Pattern of Neocirrhites armatus (Cirrhitidae) with Notes on Other Hawkfish Species." Environmental Biology of Fishes 42 (1995): 143–150.

Yagishita, N., T. Kobayashi, and T. Nakabo. "Review of Monophyly of the Kyphosidae (Sensu Nelson 1994), Inferred from the Mitochondrial ND2 Gene." Ichthyological Research 49 (2002): 103–108.

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: donaldsn@uog9.uog.edu Web site: <http://www.iucn.org/themes/ssc/sgs/sgs.htm>

Terry J. Donaldson, PhD