Dorsal fin continuous, with 10 spines and nine soft rays; anal fin with three spines and seven or eight soft rays; caudal fin in some with filament off upper lobe; six branchiostegal rays; gill membranes free from isthmus; subocular shelf and accessory subpelvic keel well developed; opisthotic (= intercalar) well developed (lost or fused in the three related families); 24 vertebrae. Nemipterids, part of the sparoid assemblage, appear to be most closely related to lethrinids.
As proposed in a 1980 study by G. D. Johnson and confirmed by Carpenter and Johnson (2002), the families Nemipteridae, Lethrinidae, Sparidae, and Centracanthidae (with the centracanthids being unresolved with respect to sparid genera), in phyletic sequence as given, form a monophyletic clade and could be placed in the superfamily Sparoidea. The study of Orrell et al. (2002) found Lethrinidae are sister to Sparidae and generally gave support to the monophyly of the Sparoidea. I do not recognize a formal superfamily Sparoidea in classification pending more study of other percoid families in hopes of presenting a comprehensive and monophyletic classification of the entire group.
Five genera, Nemipterus, Parascolopsis, Pentapodus, Scaevius, and Scolopsis, with about 64 species (e.g., Russell, 2001).
Family LETHRINIDAE (377)—emperors or emperor breams. Marine coastal; tropical, west Africa and Indo-West Pacific.
Dorsal fin continuous, with 10 spines and 9 or 10 soft rays; anal fin with three spines and 8-10 soft rays; six branchiostegal rays; no accessory subpelvic keel; reduced subocular shelf; 24 vertebrae.
Only one species occurs in the Atlantic Ocean—Lethrinus atlanticus. Some authors recognize the subfamily Lethrininae (the emperors—cheek scaleless, nine soft dorsal fin rays, subocular shelf absent) for Lethrinus and the subfamily Monotaxinae (the large-eye breams—cheeks each with at least three transverse rows of scales, 10 soft dorsal fin rays, subocular shelf present) for the other four genera; the phylogenetic validity of this arrangement, however, is uncertain. See above under family Nemipteridae, for comment on phylogenetic relationships of this family.
Five genera, Gnathodentex (1), Gymnocranius (8), Lethrinus (28), Monotaxis (1), and Wattsia (1), with about 39 species (e.g., Carpenter and Randall, 2003).
Family SPARIDAE (378)—porgies. Marine (very rarely brackish and freshwater); Atlantic, Indian, and Pacific.
Dorsal fin continuous, usually with 10-13 spines and 10-15 soft rays; anal fin with three spines and 8-14 soft rays; maxilla covered by a sheath when mouth closed; six branchiostegal rays; 24 vertebrae (10 + 14). Maximum length about 1.2 m.
The continental western Atlantic Sheepshead, Archosargus probatocephalus, which occasionally occurs in brackish water, is known to enter freshwater rarely in Florida. Four species of sparids occur in brackish water in Australia and one species of Acanthopagrus, which enters freshwater, is known to spawn in brackish water.
As noted in Orrell et al. (2002), six sparid subfamilies have been recognized (Boopsinae, Denticinae, Diplodinae, Pagellinae, Pagrinae, and Sparinae); they were not monophyletic in all their analyses. Their analysis supported a monophyletic Sparidae only with the inclusion of Spicara (included here, as traditionally, in Centracanthidae). See above under family Nemipteridae for comment on phylogenetic relationships of this family.
Thirty-three genera (e.g., Archosargus, Boops, Calamus, Chrysophrys, Dentex, Diplodus, Lagodon, Pagellus, Pagrus, Pimelepterus, Rhabdosargus, Sparus, and Stenotomus) with about 115 species (e.g., Orrell et al., 2002; Carpenter, 2003:1554-1577).
Family CENTRACANTHIDAE (Maenidae) (379)—picarel porgies. Marine; eastern Atlantic (including Mediterranean) and off South Africa.
Dorsal fin continuous, with 11-13 spines and 9-17 soft rays; anal fin with three spines and 9-16 soft rays; six branchiostegal rays; 24 vertebrae. The species of this group of planktivorous fishes have a highly protrusible upper jaw. Maximum length about 38 cm.
See above under family Nemipteridae, for comment on phylogenetic relationships of this family.
Two genera, Centracanthus (1) and Spicara (7), with eight species (references in Nelson, 1994).
Family POLYNEMIDAE (380)—threadfins. Marine and brackish water (some in rivers, especially in Borneo); all tropical and subtropical seas.
Pectoral fin divided into two sections, the upper with rays attached and the lower with 3-7 long unattached rays (but usually 14 or 15 in Polynemus (= Polistonemus multifilis)); two widely separated dorsal fins (one spiny and one soft rayed); pelvics subabdominal, with one spine and five branched rays; caudal fin deeply forked; mouth subterminal; 24 or 25 vertebrae. Maximum length 1.8 m, attained in Eleutheronema tetradactylum. About four species are known only from freshwater.
The previously assumed relationship of this family to the mugilids and sphyraenids (Nelson, 1984), all of which have widely separated dorsal fins, is no longer considered probable. It may be the sister group of sciaenids (Johnson, 1993). Much work on this group has been done since 1994 by R. M. Feltes and by H. Motomura and his coauthors.
Eight genera, Eleutheronema, Filimanus, Galeoides, Leptomelanosoma, Parapolynemus, Pentanemus, Polydactylus, and Polynemus, with 41 species (Motomura, 2004b, c).
Family SCIAENIDAE (381)—drums (croakers). Marine, brackish, and freshwater (particularly in South America); Atlantic, Indian, and Pacific.
Dorsal fin long, with a deep notch separating spinous from soft portion (rarely separate), first with 6-13 spines and second with one spine and usually 20-35 soft rays; anal fin with one or two spines (both are usually weak but the second may be large) and 6-13 soft rays; lateral line scales extending to the end of caudal fin; caudal fin slightly emarginate to rounded; upper bony
edge of opercle forked, bony flap present above gill opening; single barbel or a patch of small barbels on chin of some species; head with large cavernous canals (part of the lateral-line system); conspicuous pores on snout and lower jaw; vomer and palatine without teeth; swim bladder (rarely rudimentary in adults) usually with many branches; otoliths (sagitta at least) exceptionally large; vertebrae 24-30.
Sciaenids can produce sound by using the swim bladder as a resonating chamber. Some are important food fishes. They occur in shallow water, usually near continental regions, and are absent from islands in the mid-Indian and Pacific oceans. Several marine species enter estuaries, and about 28 species are restricted to freshwater (Atlantic drainages) in the Americas. The freshwater members are Aplodinotus grunniens, extending from southern Saskatchewan and Quebec to Guatemala, and species of Pachypops, Pachyurus, and Plagioscion (enters estuaries), in South America. K. Sasaki in 1989 revised the family, recognizing 10 subfamilies.
Bortone (2002) gives much information on various members of Cynoscion, especially C. nebulosus.
About 70 genera (e.g., Aplodinotus, Atractoscion, Atrobucca, Bairdiella, Cheilotrema, Cynoscion, Equetus, Genyonemus, Johnius, Larimus, Leiostomus, Menticirrhus, Micropogon, Micropogonias, Ophioscion, Otolithes, Otolithoides, Paranebris, Pogonias, Protosciaena, Roncador, Sciaena, Sciaenops, Seriphus, Stellifer, and Umbrina) with about 270 species (e.g., Schwarzhans, 1993; Chao et al., 2001; Chao, 2003).
Family MULLIDAE (382)—goatfishes. Marine (rarely brackish water); Atlantic, Indian, and Pacific.
Two long independently movable hyoid barbels (used in detecting food); body elongate; two widely separated dorsal fins, the first with 6-8 spines and second with one spine and 8 or 9 soft rays; soft dorsal fin shorter than anal fin; anal fin with one or two small spines and 5-8 soft rays; caudal fin forked; 24 vertebrae.
Goatfishes are important as a food fish. Many are brightly colored. Maximum length up to 60 cm.
Six genera, Mulloidichthys, Mullus, Parupeneus, Pseudupeneus, Upeneichthys, and Upeneus, with about 62 species (Golani, 2001; Kim and Nakaya, 2002; Randall and Myers, 2002; Randall, 2003).
Family PEMPHERIDAE (383)—sweepers. Marine and brackish water; western Atlantic, Indian, and Pacific.
Body compressed and deep; maxillae not reaching beyond center of eye; pre-orbital smooth; eye large, without adipose lid; one short dorsal fin, originating before middle of body, with 4-7 graduated spines and 7-12 soft rays; anal fin with two (very rarely) or three spines and 17-45 soft rays; lateral line scales usually 40-82; lateral line extending onto caudal fin; tubes of lateral line usually short and wide; gill rakers long and usually 25-31; luminescent organs in a few species; pyloric caeca 9 or 10; swim bladder absent in one species (Pempherispoeyi); 25 vertebrae (10 + 15). Maximum length about 30 cm.
Two genera, Parapriacanthus (about 5) and Pempheris (about 21), with about 26 species (e.g., Mooi, 1998).
Family GLAUCOSOMATIDAE (384)—pearl perches. Marine; eastern Indian and western Pacific (Japan to Australia).
Dorsal fin with eight graduated spines and 11-14 soft rays; anal fin with three spines and 12 soft rays; maxillae scaled; lateral line nearly straight and extending to tail; caudal fin lunate or truncate; vertebrae 25. Maximum length at about 1.2 m. This taxon, which shares similarities in the complex swim bladder/vertebral association and dorsal gill-arch elements with Pempheris, is thought to be closely related to the pempherids.
One genus, Glaucosoma, with four species (McKay, 1997).
Family LEPTOBRAMIDAE (385)—beachsalmon. Marine and brackish water (occasionally entering rivers); coasts of southern New Guinea, Queensland, and Western Australia.
Body compressed and deep; maxillae reaching far behind eye; preorbital serrate; eye relatively small, with adipose lid; one short dorsal fin behind middle of body (above anal fin), with four spines and 16-18 soft rays; anal fin with three spines and 26-30 soft rays; lateral line scales about 75-77; tubes in lateral line long and narrow; gill rakers short, usually 10. Maximum length about 35 cm. One species, Leptobrama muelleri.
Family BATHYCLUPEIDAE (386)—bathyclupeids. Marine oceanic; Indian, western Pacific, and Gulf of Mexico.
One dorsal fin in posterior half of body, without spines; anal fin long, with one spine; dorsal and anal fins covered with scales; premaxillae and maxillae bordering mouth; usually 31 vertebrae (10 + 21). One genus, Bathyclupea, with about five species.
Family MONODACTYLIDAE (387)—moonfishes (fingerfishes). Marine and brackish water (sometimes entering freshwater); western Africa and Indo-Pacific.
Body strongly compressed and deep (deeper than long in some); pelvic fins present in juveniles but absent or vestigial in adults of Monodactylus; dorsal fin single and with a long base, covered with scales and 5-8 short graduated spines; anal fin with three spines, long base; scales cycloid or ctenoid.
Moonfishes are occasionally sold as aquarium fishes. They are often of a silvery color. The three species of Monodactylus often ascend rivers, and some populations may live in freshwater. For the possible relationships of this and the next several families see the comment on Squamipennes under Chaetodontidae.
Two genera, Monodactylus and Schuettea, with about five species. Some authors place Schuettea (with two species in New South Wales and Western Australia) in the Monodactylidae, but Y. Tominaga in 1968 recommended placement in a family of its own. It differs from other monodactylids in a few characters (e.g., normally developed pelvic fins, cycloid scales, teeth absent from endopterygoid and ectopterygoid) and is provisionally retained in the family as a conservative measure.
Family TOXOTIDAE (388)—archerfishes. Marine coastal, brackish, and freshwater; from India to Philippines and Australia and Polynesia.
Body deep and compressed, greatest body depth 1.8-2.5 times in standard length; eye large; dorsal fin with 4-6 strong spines and 11-14 soft rays; anal fin with three spines and 15-18 soft rays; length of soft dorsal much shorter than soft portion of anal; mouth large, terminal (lower jaw protruding), and highly protractile; lateral line scales about 25-47; seven branchiostegal rays; 24 vertebrae (10+14).
Archerfishes are capable of forcefully ejecting squirts of water from their mouths and downing insects. The widespread Toxotes jaculator, extending from India to New Hebrides, is normally found in brackish water near mangroves, while the others frequently occur in freshwater (often well inland). Maximum length 40 cm, attained in T. chatereus; usually under 16 cm. One genus, Toxotes, with six species (reviewed by G. R. Allen in 1978).
Family ARRIPIDAE (389)—Australasian salmon (kahawai). Marine; South Pacific (southern Australia to New Zealand region).
Dorsal fin with 9 spines and 13-19 soft rays; anal fin with three spines and 9 or 10 soft rays; gill membranes free from isthmus; anal fin much shorter than the soft dorsal; caudal fin forked; 25 vertebrae. Maximum length about 90 cm. One genus, Arripis, with four species (Paulin, 1993).
Family DICHISTIIDAE (Coracinidae) (390)—galjoen fishes. Marine coastal and brackish water; South Africa and Madagascar.
Body relatively deep; mouth small; dorsal fin with 10 spines and usually 18-23 soft rays; anal fin with three spines and usually 13 or 14 soft rays; gill membranes fused with isthmus; some teeth incisiform.
This inshore fish is a highly sought after sports fish in South Africa (Smith and Heemstra, 1986).
This family was recognized under the name Coracinidae in Nelson (1994), with the generic name Coracinus; Eschmeyer (1998) noted that Coracinus was published in a rejected work and is not available (the family name should be after the next available synonym).
Although monophyly of this family could not be confirmed, Leis and van der Lingen (1997) found larval evidence to support the historical linking of species of Microcanthinae, Scorpidinae, Girellinae, and Kyphosinae (recognized by them at the family level; a grouping in the earlier study of G. D. Johnson) with possibly some affinity to the Arripidae (but not to the Ephippidae or Drepaneidae).
One genus, Dichistius, with about two species (e.g., Leis and van der Lingen, 1997).
Family KYPHOSIDAE (391)—sea chubs. Marine; Atlantic, Indian, and Pacific.
Three spines and 10-28 soft rays; dorsal fin with 9-16 spines and 11-28 soft rays; 24-28 vertebrae (34 in Graus). Members of the first two subfamilies, except for Graus, are herbivorous, primarily consuming algae, while the others are primarily carnivorous. All are usually found near shore.
Monophyly for the group has not been established, and the following subfamilies, or combinations thereof, are often recognized as separate families (it may be desirable to recognize the subfamily Parascorpidinae at the family level). G. D. Johnson and R. A. Fritzsche in a 1989 study provided evidence for the monophyly of a taxon including the first three subfamilies (which they rank as families, as did the 1984 study of G. D. Johnson). Yagishita et al. (2002) made a valuable start in determining whether or not this family is monophyletic, but as they stated, many more taxa need to be included in an analysis. Their results suggested that Scorpis, Labracoglossa (these two being sister taxa), Girella, Microcanthus, and Kyphosus are part of a monophyletic group but one shared with Kuhlia and Oplegnathus. Much of the basis for the generic composition of these taxa is based on the G. D. Johnson studies. See comments above under family Dichistiidae. Sixteen genera with about 45 species.
Subfamily Girellinae (Nibblers). Some incisiform teeth present; maxilla concealed beneath suborbital bone. Pacific (primarily Philippines to Australia but extending to California where the Opaleye, Girella nigricans, is a common inshore species) with G. zonata in the Atlantic. Graus occurs off Chile.
Two genera, Girella and Graus, with about 17 species (e.g., Yagishita and Nakabo, 2000).
Subfamily Kyphosinae (Rudderfishes) . Some incisiform teeth; maxilla exposed. Atlantic, indian, and Pacific.
Four genera, Hermosilla, Kyphosus, Neoscorpis, and Sectator (e.g., Sakai and Nakabo, 2004) with 13 species.
Subfamily Scorpidinae (Halfmoons). No incisiform teeth; pelvics well behind pectorals. indo-Pacific (to California).
Four genera Bathystethus, Labracoglossa, Medialuna, and Scorpis, with about seven species.
Subfamily Microcanthinae. Recognized as a family by Johnson (1984:469).
Five genera, Atypichthys, Microcanthus, Neatypus, and Tilodon; Vinculum may also belong here. About seven species.
Subfamily Parascorpidinae (Jutjaws) . Mouth large with lower jaw projecting forward; upper jaw not protractile; 27 vertebrae. Maximum length 60 cm.
It may be desirable to recognize this subfamily at the family level; its affinities are uncertain.
One species, Parascorpis typus, known only from South Africa (e.g., Smith and Heemstra, 1986).
Family DREPANEIDAE (392)—sicklefishes. Marine; Indo-West Pacific and West Africa.
Dorsal fin with 13 or 14 spines and 19-22 soft rays; anal fin with three spines and 17-19 soft rays; mouth markedly protractile; pectoral fins longer than head, falcate; maxilla distally exposed; subocular shelf absent; 24 vertebrae. The recognition of this family for Drepene follows the 1984 study of G. D. Johnson, who placed it next to the Dichistiidae (= Coracinidae).
The spelling of this family name in previous editions was Drepanidae. As noted in Eschmeyer, 1998, Vol 3:2889, Opinion 1046 of the ICZN required that it be Drepaneidae. See below under the suborder Acanthuroidei for evidence from Tang et al. (1999) that it belongs within that suborder. One genus, Drepane, with two or three species.
Family CHAETODONTIDAE (393)—butterflyfishes. Marine; tropical to temperate Atlantic, Indian, and Pacific (primarily tropical Indo-West Pacific).
Body strongly compressed; no spine at angle of preopercle (small serrations may be present on the preopercle); well-developed pelvic axillary process; head region in larval (tholichthys) stage of most species covered with bony plates; dorsal fin continuous or with slight notch, with 6-16 spines and 15-30 soft rays, no procumbent spine; anal fin with 3-5 (usually 3) spines and 14-23
soft rays; caudal fin with 15 branched rays (17 principal), margin rounded to emarginate; scales extending onto the dorsal and anal fins; mouth small, terminal, protractile (the two species of the Indo-Pacific Forcipiger have a very elongate snout); gut coiled many times; swim bladder with two anteriorly directed processes; 24 vertebrae (11 + 13).
Most species of butterflyfish have brightly colored patterns. Also, most have a dark band running across the eye, and many have an "eyespot" on the dorsal or posterior part of the body—both patterns may serve to confuse predators. Butterflyfishes generally occur near coral reefs and at depths of less than 20 m, but a few go to at least 200 m. A few species occur in brackish water. Most species are in the Australian to Taiwan region. Only 13 species occur in the Atlantic and 4 in the eastern Pacific. Butterflyfishes are known to feed on coral polyps (but are not known to break off coral) and on other invertebrates.
G. Cuvier included chaetodontids and all other fishes with the proximal portion of the dorsal and anal fins covered with scales in the family Squamipinnes (= Squamipennes). Various authors subsequently employed this taxon, usually at the subordinal level following Jordan and Evermann's 1898 work, with varying membership. Nelson (1994) noted the families forming the squamipinnes group and referred to the 1983 study of H.-K. Mok and S.-C. Shen, who provided evidence that the squamipinnes, the acanthuroids, and the tetraodontiforms form a monophyletic taxon.
Butterflyfishes and angelfishes were, until the mid-1970s, combined in the same family; however, W. E. Burgess in a 1974 study gave reasons for recognizing them in separate families (and noted many morphological differences, including those in osteology, between the two groups), and Bellwood et al. (2004) confirmed that the families are monophyletic (but not necessarily each other's closest relatives).
Eleven genera (e.g., Chaetodon, Chelmon, Coradion, Forcipiger, Hemitaurichthys, Heniochus, Johnrandallia, and Prognathodes) with about 122 species (Allen et al., 1998; Kuiter and Debelius, 1999; Burgess, 2001; W. L. Smith et al., 2003; Burgess, 2003).
Family POMACANTHIDAE (394)—angelfishes. Marine; tropical Atlantic, Indian, and Pacific (primarily in western Pacific).
Body strongly compressed; strong spine at angle of preopercle; no well-developed pelvic axillary process; larval stage lacking bony head plates but having spiny scales; dorsal fin continuous, with 9-15 spines and 15-37 soft rays, no procumbent spine; anal fin with three spines and 14-25 soft rays; dorsal and anal fins with elongate extension on hind margin in many species (shown in figure); caudal fin with 15 branched rays, margin rounded to lunate (strongly lunate, often with produced lobes, in some species of Genicanthus); swim bladder lacking anteriorly directed processes; 24 vertebrae (10 + 14).
Angelfishes have striking color patterns and in many species the pattern in juveniles differs markedly from that of adults. They generally occur near coral reefs at depths of less than 20 m (very seldom below 50 m).
Eight genera, Apolemichthys, Centropyge (synonym Sumireyakko), Chaetodontoplus, Genicanthus, Holacanthus, Paracentropyge, Pomacanthus, and Pygoplites, with about 82 species (e.g., Pyle, 1997; Allen et al., 1998; Allen and Steene, 2004; Randall and Carlson, 2000; Burgess, 2003; Bellwood et al., 2004).
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