Genus EPINEPHELUS BLOCH, 1793

Epinephelus BLOCH, 1793: 11 (type species, Epinephelus marginalis BLOCH, 1793 [= E. fasciatus (FORSSKAL, 1775)], designated under the plenary powers of the International Commission on Zoological Nomenclature, Opinion 93).

Merou BONAPARTE, 1831: 167 (type species, Perca gigas BRÜNNICH, 1768 [nomen dubium] by subsequent designation of JORDAN, 1919: 175).

Cerna BONAPARTE, 1833: puntàta 10 (type species, Perca gigas BRÜNNICH by monotypy).

Cynichthys SWAINSON, 1839: 168, 201 (type species, C. flavo-purpuratus [= Perca flavapurpurea BENNETT, 1830 = E. flavocaeruleus (LACEPÈDE, 1802)]).

Cernua COSTA, 1849: 1 (not available; unjustified emendation of Cerna BONAPARTE,‘ preoccupied by Cernua FLEMING, 1828: 212 (a genus of percid fish).

Hyporthodus GILL, 1861: 98 (type species Hyporthodus flavicauda GILL [= E. niveatus VALENCIENNES, 1828] by monotypy.

Schistorus GILL, 1862: 236 (type species, Serranus mystacinus POEY, 1852 by monotypy).

Labroperca GILL, 1862: 236 (type species, Serranus labriformis JENYNS, 1843 by monotypy).

Promicrops POEY, 1868: 287 (type species, Serranus Guasa POEY, 1861 [= E. itajara (LICHTENSTEIN, 1822)] by monotypy; genus attributed to GILL by POEY, but the diagnosis is POEY's).

Priacanthichthys DAY, 1868: 193 (type species, Priacanthichthys maderaspatensis DAY, 1868 [= E. latifasciatus (TEMMINCK & SCHLEGEL, 1842)] by monotypy).

Merus POEY, 1874: 39 (type species, Epinephelus marginalis BLOCH; proposed as a replacement name for Epinephelus BLOCH).

Homalogrystes ALLEYNE & MACLEAY, 1877: 268 (type species, Homalogrystex guntheri ALLEYNE & MACLEAY, 1877 [= E. coioides (HAMILTON, 1822)] by monotypy).

Itaiara VAILLANT& BOCOURT, 1878: 70 (type species, Serranus itajara LICHTENSTEIN, by monotypy).

Hyposerranus KLUNZINGER, 1884: 3 (type species, Serranus morrhua VALENCIENNES, 1833 by subsequent designation of JORDAN, 1920; proposed as a subgenus of Serranus).

Phrynotitan GILL, 1885: 225 (type species, Batrachus gigas GÜNTHER, [= E. lancolatus] by monotypy).

Garrupa JORDAN, in JORDAN & EIGENMANN, 1890: 350, 353 (type species, Serranus nigritus HOLBROOK, 1855 by original designation; proposed as a subgenus of Epinephelus).

Enneistus JORDAN & EVERMANN, 1896: 1147 (type species, Bodianus acanthistius GILBERT, 1892 by monotypy; proposed as a subgenus of Bodianus).

Stereolepoides FOWLER, 1923: 382 (type species, Stereolepoides thompsoni FOWLER, 1923 [= E. lanceolatus] by monotypy.

Vivero JORDAN & EVERMANN, 1927: 505 (type species, Serranus morio VALENCIENNES, 1828 by monotypy; proposed as a subgenus of Epinephelus).

Serrihastaperca FOWLER, 1944: 384 (type species, Serrihastaperca exsul FOWLER, 1944 by original designation).

Altiserranus WHITLEY, 1947: 150 (type-species Serranus jayakari BOULENGER, 1889 [= E, multinotatus (PETERS, 1876)] by original designation).

Diagnosis: Body elongate, robust (subcylindrical), oblong or deep and compressed; body depth greater than, subequal to or less than head length and contained 2.3-3.7 times in SL, the body width 1.8-2.8 in the depth; head length 2.1 -2.8 in SL; preorbital depth 6.7- 15 times in head length; preopercle rounded or angular, the posterior edge serrate, with the serrae at the angle more or less enlarged; a few species with small serrae (mostly covered by skin) on the ventral edge. Dorsal fin usually with 11 spines (10 spines in E. analogus, exsul and nigritus, 9 in E. acanthistius) and 12- 19 rays; length of base of soft-rayed part of dorsal fin not more than base of spinous part. Anal fin with 3 distinct spines and 7- 10 (very rarely 7 or 10) rays. Caudal fin rounded, truncate or concave, with 8 + 7 branched rays and 8- 10 + 7-10 procurrent rays. Pectoral fin rounded, with the middle rays longest. Scales on body ctenoid or smooth. Canines present at front of jaws, but they may be small in some species; no distinctly enlarged canine tooth at midside of lower jaw; teeth present on palatines; maxilla of adults without a distinct bony knob on ventroposterior corner, but there may be an abrupt step or hook-like process (covered by the upper lip) on the distal part of the ventral edge; supramaxilla well developed.

Supraneural bones 2; dorsal and anal fins without trisegmental pterygiophores; rear edge of first dorsal pterygiophore with or without excavation for tip of 2nd neural spine; epipleural ribs on first 8- 10 vertebrae. The diversity of cranial morphology of the many species assigned to Epinephelus makes it difficult to recognize diagnostic cranial characters for the genus.

GEOGRAPHICAL DISTRIBUTION

The genus Epinephelus is represented in tropical and subtropical latitudes of all three major oceans, as well as in the Mediterranean Sea.

REMARKS

C. L. SMITH (1971) demoted the genus Promicrops (comprising E. itajara and E. lanceolatus of the Indo-West Pacific) to a subgenus of Epinephelus and stated that these two species “are highly specialized and distinctive although their alliance with other species of Epinephelus is clear". As justification for his recognition of Promicrops as a subgenus, SMITH (1971: 152) mentioned some distinctive features of the cranium of E. itajara, especially "a heavy strut of bone some connecting the posterior face of each lateral ethmoid with the shaft of the parasphenoid (fig. 17).“ This posterior ethmoid strut is not known for any other species of grouper.

E. itajara and E. lanceolatus also differ from most other species of the genus in having the tubes of the lateral-line scales with 4-6 radiating branches. Except for large adults of E. malabaricus and E. coioides (which have a few anterior lateral-lure scales With branched tubules), the lateral-line scales of other species of Epinephelus have unbranched tubes.

Epinephelus is compared with the genus Cephalopholis in the account of that genus (above). The relationships of the genus Epinephelus are obscure, because this taxon (like other grouper genera) is defined by superficial (not clearly synapomorphic) characters that may or may not indicate the congeneric status of the included species. Nevertheless, the genus can easily be separated from the other eastern Atlantic genera, as indicated in the key to species (above). A comparison of Epinephelus with the grouper genera that are not represented in the eastern Atlantic is given in the papers of RANDALL and HEEMSTRA (in press) and HEEMSTRA and RANDALL (in press).

The genus Epinephelus, as here defined, comprises some 97 species and is thus the most speciose genus of serranid fishes. It is well represented in the tropical and subtropical waters of all three major oceans. Most species (67) are found in the vast Indo-West Pacific region (see RANDALL and HEEMSTRA, in press, for a revision of the Indo-West Pacific species). Eight species occur in the eastern Pacific; 11 are known from the western Atlantic, and 9 species are found in the eastern Atlantic and Mediterranean.

Epinephelus species are generally found on coral or rocky reefs, but a few species (e.g., E. aeneus) are commonly taken with trawls over sandy/mud bottoms. Some species occur in deep water (to at least 525 m.), but most are found in depths of 10- 200 m. The two largest species ( E. itajara and E. lanceolatus), which grow to well over 2 metres in length and a weight of over 400 kg, are often found in estuaries and harbours.

The reproduction of a few species has been studied, and they appear to be protogynous hermaphrodites; but the picture is complicated in some species by the occurence of males that are much smaller than some females. It may be that not all females change sex, and perhaps some males do not go through a previous female stage.