Family Centrarchidae 349sunfishes Freshwater North America

Fin Spines

Suborbital bones present in addition to the lachrymal; dentary and angular penetrated by lateral line; lateral line present on body, sometimes incomplete; anal fin spines usually three (or fewer, typically in species of Enneacanthus, Lepomis, and Micropterus) or five (or more, typically in species of Acantharchus, Ambloplites, Archoplites, Centrarchus, and Pomoxis); dorsal fin usually with 5-13 spines (most with 10-12); pseudobranch small and concealed; branchiostegal rays 6 or 7; gill membranes separate; vertebrae 28-33.

Most sunfishes are nest builders. The male hollows out a small depression with his tail and then guards the eggs. Centrarchids are an important sports fish and have been introduced into many areas beyond their native range. Some, such as Lepomis macrochirus, the Bluegill, have been used in physiological and ecological experimental work. Maximum length about 83 cm, attained in Micropterus salmoides (Largemouth Bass).

Centrarchids were widespread west of the Rocky Mountains by the Late Miocene; the fossil record includes Plioplarchus and species of extant genera.

Eight genera with 31 species. The classification with genera recognized is based on Roe et al. (2002) and the species numbers are from Nelson et al. (2004). Gilbert (1998) gives a type catalogue of recent and fossil taxa.

Subfamily Centrarchinae. Five genera, Ambhplites (4), Archoplites (1, the Sacramento Perch, is the only living centrarchid native west of the Rocky Mountains), Centrarchus (1), Enneacanthus (3), and Pomoxis (2, crappies), with 11 species.

Subfamily Lepominae. One genus, Lepomis (12, upper figure, synonym Chaenobryttus).

Subfamily uncertain. The genera Acantharchus (1) and Micropterus (7, basses) (lower figure) are of uncertain relationships and not placed in either subfamily until their relationships are better known.

Family PERCIDAE (350)—perches. Freshwater; Northern Hemisphere.

Two dorsal fins, separate or narrowlyjoined (broadlyjoined in Zingel); one or, usually, two anal spines (the second is usually weak); pelvic fins thoracic, with one spine and five soft rays; premaxilla protractile or nonprotractile; bran-chiostegal rays 5-8; branchiostegal membrane not joined to isthmus (may be united to each other or not); pseudobranchiae well developed to rudimentary; no subocular shelf; supramaxilla absent; one or no predorsal bones (rarely two) (interneural before first pterygiophore); vertebrae 32-50. Maximum size up to 90 cm, attained in Sander vitreus (Walleye); most species much smaller.

Ten genera with 201 species (187 in North America and 14 in Eurasia). The number of species in the genera endemic to North America is from Nelson et al. (2004), which is based on a series of continuing studies describing new species. The classification of Song et al. (1998) is followed below where three monophyletic groups are each recognized as subfamilies. The two tribes of the subfamily Percinae in Nelson (1994) are now recognized as subfamilies (i.e., Percinae and Etheostomatinae). Mioplosus, an Eocene Green River Formation percoid, may have its closest affinity with the percids (Grande, 1984), but more work is needed before we can understand its relationships (Grande, 2001).

Subfamily Percinae. Anteriormost interhaemal bone greatly enlarged; anal spines usually well developed; preopercle strongly serrate; usually seven or eight branchiostegal rays; body compressed; anal spines prominent; swim bladder well developed.

Three genera with eight species: the circumpolar Perca (3, the Eurasian P. fluviatilis which has been introduced into South Africa, Australia, and New Zealand, the almost identical North American P. flavescens, and P. schrenki of the Balkhash and Alakul' lakes area of Asia); the European and western Asian Gymnocephalus (4); and Percarina demidoffi of the northern Black Sea area. The biology of various species, especially of Perca, is given by Craig (2000).

Subfamily Luciopercinae. Anteriormost interhaemal no larger than posterior ones; anal spines weak; lateral line extending onto tail.

Three genera and nine species. Contains the genus of predaceous pikeperches, Sander (synonym Stizostedion), possessing a well-developed swim bladder, with three species in Europe (including the Caspian and Aral seas) and two species (Sauger and Walleye) in North America. It also contains two genera of European darterlike fishes lacking a swim bladder, Zingel, with three species, of the Danube, Rhone, and Vardar systems and the very restricted Romanichthys valsanicola of Romania.

Subfamily Etheostomatinae. Anteriormost interhaemal bone greatly enlarged; anal spines usually well developed; preopercle margin smooth or

partly serrated; usually five or six branchiostegal rays; body slightly compressed or fusiform; anal spines moderately prominent; swim bladder reduced or absent. Seldom over 11 cm.

Body Compressed Fusiform

Four genera of North American darters: Ammocrypta (6), Crystallaria (with one species, C. asprella), Etheostoma (136), and Percina (41), giving a total of 184 described species (species listed in Nelson et al., 2004). There are many studies on North American darters, with many new species described in the last few decades (as noted in Nelson et al., 2004); much of this work has been done by L. M. Page and his students (e.g., Burr and Page, 1993; Page et al., 2003).

Family PRIACANTHIDAE (351)—bigeyes (catalufas). Marine; tropical and subtropical, Atlantic, Indian, and Pacific.

The Fishes Indian Pacific

Eyes very large; mouth large, strongly oblique; dorsal fin continuous, usually with 10 spines and 11-15 soft rays; anal fin with three spines and 10-16 soft rays; caudal fin with 16 principal rays (14 branched), slightly emarginate to rounded; membrane present connecting the inner rays of the pelvic fin to the body; scales modified spinous cycloid (with strong spines but not ctenoid); scales on the branchiostegal membrane; color usually bright red; vertebrae 23 (very few other percoids have so few vertebrae).

Bigeyes are usually carnivorous and nocturnal. The tapetum lucidum, a brilliant reflective layer producing "eyeshine," may be of a unique form among teleosts. Maximum length about 65 cm TL.

Four genera, Cookeolus, Heteropriacanthus, Priacanthus, and Pristigenys, with about 18 species (e.g., Starnes, 2003). The fossil record is discussed in the 1988 revision of W. C. Starnes.

Family APOGONIDAE (352)—cardinalfishes. Marine, some brackish water, a few in streams in the tropical Pacific; Atlantic, Indian, and Pacific.

Two separated dorsal fins, the first with 6-8 spines and the second with one spine and 8-14 soft rays (Paxton has a continuous dorsal fin); anal fin with two spines and 8-18 soft rays; scales usually ctenoid, but cycloid in several groups and absent in Gymnapogon; seven branchiostegal rays; usually 24 vertebrae (10 + 14). Several other families have widely separated dorsal fins, but this is the only one in which the distal radial of the last spine is short (vs elongate) (Johnson, 1993). Species of Siphamia have a ventral luminous organ. Many of the species are mouthbreeders; it is suspected that in some only the males incubate the eggs, whereas in others it is only the females. Most species nocturnal. Maximum length usually about 20 cm, and most are less than 10 cm.

The nine species of Glossamia are found only in freshwater; they occur primarily in New Guinea but also in Australia. A few species of Apogon occur in estuaries and the lower reaches of rivers. The relatively deep dwelling (60-290 m) Apogon gularis is unique among apogonids in having the anus located just behind the origin of the pelvic fins. Baldwin and Johnson (1999) supported the recognition of the following two subfamilies.

About 23 genera and roughly 273 species.

Subfamily Apogoninae. Oral incubation of eggs in at least many species. About 19 genera (e.g., Apogon, Apogonichthys, Archamia, Astrapogon, Cercamia, Cheilodipterus, Coranthus, Foa, Fowleria, Glossamia, Neamia, Phaeoptyx, Rhabdamia, Siphamia, and Vincentia), with about 260 species (Fraser, 2000, 2005; Allen, 2001; Greenfield, 2001; Gon, 2003; Gon and Randall, 2003a, b).

Subfamily Pseudaminae. Large canine teeth always present on dentary and premaxillae; lateral line incomplete or absent; scales cycloid or absent. Four genera, Gymnapogon, Paxton, Pseudamia, and Pseudamiops, with roughly 13 species (e.g., Baldwin and Johnson, 1999).

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